CIHM 
Microfiche 
Series 
(IVIonographs) 


ICIMH 

Collection  de 
microfiches 
(monographies) 


Canadian  Institute  for  Historical  Microreproductions  /  Institut  Canadian  da  microraproductions  historiquas 


Tech 

The  Institute  h£ 
copy  available  I 
may  be  bibliogr 
the  images  ii 
significantly  ch 
checked  below. 


n 


D 


Coloured  i 
Couvertur 

Covers  da 
Couvertur 

Covers  re! 
Couvertur 

Cover  title 

Coloured  i 


Coloured  i 
Encro  de  < 


r~yT   Coloured} 


D 
D 
D 


D 


Q 


Planches  i 

Bound  witi 
Reli6  avec 

Only  editio 
Seule  ^diti 

Tight  bindii 
interior  mj 
I'ombre  oi 
int^rieure. 

Blank  leavi 
within  the  ti 
omitted  fro 
blanches 
apparaissE 
possible,  c 

Additional 
Commenta 


This  item  is  lilmed  t 
Ce  document  est  fill 

lOx 


12x 


chnical  and  Bibliographic  Notes  /  Notes  techniques  et  bibliographiques 


!  has  attempted  to  obtain  the  best  original 
le  for  filming.  Features  of  this  copy  which 
^graphically  unique,  which  may  alter  any  of 
i  in  the  reproduction,  or  which  may 
change  the  usual  method  of  filming  are 

)W. 

ed  covers  / 
lure  de  couleur 

I  damaged  / 

lure  endommag^e 

;  restored  and/or  laminated  / 
lure  restaur^e  et/ou  pellicul^e 

title  missing  /  Le  titre  de  couverture  manque 

3d  maps  /  Cartes  gdographiques  en  couleur 

3d  ink  (i.e.  other  than  blue  or  black)  / 
Je  couleur  (i.e.  autre  que  bieue  ou  noire) 

»d  plates  and/or  illustrations  / 
es  et/ou  illustrations  en  couleur 

with  other  material  / 
vec  d'autres  documents 

jition  available  / 
Idition  disponible 

nding  may  cause  shadows  or  distortion  along 
margin  /  La  reliure  serree  peut  causer  de 
!  ou  de  la  distorsion  le  long  de  la  marge 
re. 

javes  added  during  restorations  may  appear 
le  text.  Whenever  possible,  these  have  been 
from  filming  /  Use  peut  que  certaines  pages 
es  ajout6es  lors  d'une  restauration 
ssent  dans  le  texte,  mais,  lorsque  cela  etait 
3,  ces  pages  n'ont  pas  et6  fiim^es. 


L'Institut  a  microfilmd  le  meilleur  exemplaire  qu'il  lui  a 
m  possible  de  se  procurer.  Les  details  de  cet  exem- 
plaire qui  son!  peut-6tre  uniques  du  point  de  vue  bibli- 
ographique.  qui  peuvent  modifier  une  image  reproduite, 
ou  qui  peuvent  exiger  une  modification  dans  la  m^tho- 
de  normale  de  filmage  son!  indiqu^s  ci-dessous. 

I     I  Coloured  pages  /  Pages  de  couleur 

I I  Pages  damaged  /  Pages  endommag6es 


D 


Pages  restored  and/or  laminated  / 
Pages  restaurdes  et/ou  pellicul^es 


Q  Pages  discoloured,  stained  or  foxed  / 
Pages  d^colordes,  tachet^es  ou  piqu^es 

Pages  detached  /  Pages  d6tach6es 

\yA   Showthrough  / Transparence 

I      I  Quality  of  print  varies  / 


D 
D 


D 


Quality  in^gale  de  I'impression 

Includes  supplementary  material  / 
Comprend  drj  materiel  suppl^mentaire 

Pages  wholly  or  partially  obscured  by  errata  slips, 
tissues,  etc.,  have  been  refilmed  to  ensure  the  best 
possible  image  /  Les  pages  totalement  ou 
partiellement  obscurcies  par  un  feuillet  d'errata,  une 
pelure,  etc.,  ont  et6  fiim^es  a  nouveau  de  fa^on  a 
obtenir  la  meilleure  image  possible. 

Opposing  pages  with  varying  colouration  or 
discolourations  are  filmed  twice  to  ensure  the  best 
possible  image  /  Les  pages  s'opposant  ayant  des 
colorations  variables  ou  des  decolorations  sont 
filmdes  deux  fois  afin  d'obtenir  la  meilleure  image 
possible. 


idl  comments  / 

mtaires  suppl^mentaires: 


Various  pagings. 


ed  at  the  reduction  ratio  checked  below  / 

A  filme  au  taux  de  reduction  indique  ci-dessous. 


14x 

18x 

22x 

26x 

30x 

7 



16x 


20x 


24x 


28x 


32x 


The  copy  filmed  hare  has  baan  raproducad  thanks 
to  the  ganarosity  of: 

Engineering  Sciences  Library, 
Queen's  University 


L'axamplaira  filmA  fut  raproduit  grice  i  la 
gAn^rositA  da: 

Engineering  Sciences  Library, 
Queen's  University 


Tha  imagas  appearing  hare  are  the  bast  quality 
possible  considering  the  condition  and  legibility 
of  the  original  copy  and  in  keeping  with  the 
filming  contract  specifications. 


Original  copies  in  printed  paper  covers  are  filmed 
beginning  with  the  front  cover  and  ending  on 
the  last  page  with  a  printed  or  illustrated  impres- 
sion, or  the  back  cover  when  appropriate.  All 
other  original  copies  are  filmed  beginning  on  the 
first  page  with  a  printed  or  illustrated  impres- 
sion, and  ending  on  the  last  page  with  a  printed 
or  illustrated  impression. 


The  last  recorded  frame  on  each  microfiche 
shall  contain  tha  symbol  —»>  (meaning  "CON- 
TINUED"), or  tha  symbol  V  (meaning  "END"), 
whichever  applies. 

Maps,  plates,  charts,  etc.,  may  be  filmed  at 
different  reduction  ratios.  Those  too  large  to  be 
entirely  included  in  one  exposure  »f  filmed 
beginning  in  the  upper  left  hand  corner,  left  to 
right  and  top  to  bottom,  as  many  frames  as 
required.  The  following  diagrams  illustrate  the 
method: 


Les  imagas  suivantas  ont  iti  raproduites  avec  la 
plus  grand  soin.  compte  tenu  da  la  condition  at 
de  la  nettetA  de  Taxemplaira  filmA.  et  Bn 
conformity  avec  las  conditions  du  contrat  de 
filmaga. 

Les  exemplairas  originaux  dont  la  couvertura  en 
papier  est  imprimis  sont  filmAs  en  commencant 
par  la  premier  plat  at  en  terminant  soit  par  la 
derniAre  page  qui  comporte  unc  ampreinte 
d'impression  ou  d'illustration.  soit  par  le  second 
plat,  salon  le  cas.  Tous  les  autres  exemplairas 
originaux  sont  filmis  en  commencant  par  la 
premiere  page  qui  comporte  une  empreinte 
d'impression  ou  d'illustration  at  an  terminant  par 
la  darniire  page  qui  comporte  une  telle 
ampreinte. 

Un  des  symboles  suivants  apparaltra  sur  la 
derniire  image  de  cheque  microfiche,  selon  le 
cas:  la  symbols  — ^>  signifie  "A  SUIVRE ',  le 
symbols  V  signifie  "FIN". 

Les  cartas,  planches,  tableaux,  etc.,  peuvent  iue 
filmAs  A  des  taux  de  reduction  diffirents. 
Larsque  le  document  est  trop  grand  pour  fttre 
raproduit  en  un  seul  cliche,  il  est  filmi  i  partir 
de  Tangle  supirieur  gauche,  de  gauche  i  droite, 
et  de  haut  en  bas,  en  prenant  le  nombre 
d'images  nicessaire.  Les  diagrammes  suivants 
illustrent  la  mithode. 


1 

2 

3 

1 

2 

3 

4 

5 

6 

MIC»OCOrr    RfSOLUTION    TIST   CHART 

ANSI  and  ISO  lESI  CHART  No    ? 


1.25 


!fia  Ilia 

It  iii    '^ 

^    i^     ill  21 


u 


1.8 


1.6 


A     APPLIED  IM^GE    Inc 


-  ''6:    -Si'       IJ300    ■    Phor 


PHYSIOLOGY  AND  BIOCHEMISTRY 
IN  MODERN  MEDICINE 


I'.V 


.1.  .1.   I.'.  MACLKOI),  M.n. 


lliOFKSSDK    UK   I'llNSKM.IX.I     IN    TllK    I   M\  KKMTV    OK   TUKdNTO,    TOKONTO,    CVNAIIA;    KORMKKI.Y 

nMiKKssui;  oi    ni\.si()i.iM,\    i.\    juk   uk.stkkn    iuskuvk   inivkiisity. 

fl.F.VFI.AN'n,   OHIO 


AssisTii,  iiv    K,,v  (.'.  I'lAiic  i;.   It, A..  .M.I). 

I>ir.,ti.i     (.t     th.'    i',ir.liiMcs|iiint"ry    l.jilioratoiy     „r     l.:ik.'>i.|r     lli.-i|.it;il. 
•  •l.'Mhiii.l,  Ohio 


.\M)  I'.V   uTIIKliS 


U777/  •'.;.,■  //./,r.v77,M770.V.v.  I.XC/.I  /n.\<! 

n  i'i..\ri:s  is  coLOh-s 


ST.   I.OII.s 
•  '.  \'.  MOSMV  CO.MI'A.W 

liMS 


0  f':-"i 
•  Mil? 


I'Ms.    1^ 


Mi^Bs    I     \trA\s 


S'    I.  on  is 


TO 
.M.   W.   M. 


550*) 


)  I 


I'KKI  A(  K 


'I'lif  ii(rc»il.v    uf  alliillintf  tlio  vatiiiii>  siilijicis  oi'  llic  iiii'iliuul  fiirric- 

ilhllil    III    ililTi'lciil     |ii'liii(|s,    ,11    tllill    tlic    liiiiir    NitiiMlv    Mitillllii'    >llli,J4rt> 

all-  (■.iiii|ili't<'<l  ill  llu'  carliff  viais,  lia^  llic'  ureal  <liHa<lvaiita({<'  that  tin' 
-tiKlciit,  liriiiu  till  luiiyi'i-  in  tiiiicli  with  hilmiiitui  \  «iiik,  fails  to  t'iii|)liiy 
till'  scicntidf  Iiii()\\  l('i1i;t'  uilli  full  ,ii|\  nnlau'''  in  'Ih'  sniiitiini  of  his  clin- 
ical pioliliiiis.  Ill'  iN  apt  til  ii'LTiiiil  till'  lir^I  ivMi  iir  three  \eais  in  the 
laliiiratiil.  ilepai  I  iinlits  as  iMeiiiiM'i|iieiil  iai  in  eiiiii|)aliMi|i  \\ith  the  Mlp- 
|MiM'(ll\  iiliile  pliietiiMJ  ilisti'lietiiili  nlVeleil  iJHlini.'  tlii'  >l|liM'i|lleill  elillieai 
vears.      ife  is  taiiuht   li.\    his  ialinialiiiv    ilist  riii  tm  s  to  uliseiAe  aeeiiialelv, 

ami  ti)  eiiiiilate  tl Iisei\ei|    fai's,  s(i  Ijiat   lie  may  I iialileil  tn  draw 

i-ii|iell,siiiM>  as  III  the  hiiilil.cr  iif  wiifkiliv  iif  the  \aiiiills  fllheliiilis  iif  tile 
animal  linily  in  health,  ainl  licl'me  priieeeiliiijr  to  his  eliiiieal  studies,  he 
is  re(|uired  I"  shin\  a  plnlieieiiey  in  seieiltide  Uliiiw  ledu'e,  lieeailse  it  js 
I'eeiiL'lii/ed  that  this  must  seixe  as  the  liiisis  iipiiM  whii'li  liis  knuw  ledue 
'if  disease  is  tn  lie  liuitt.  Whi-M  the  eliiiie  is  leaehed,  hnwevef,  the  nieth 
mis  iif  the  scientist  iiie  lint  ill f iei|in  lit ly  east  aside  and  an  uiidei'staiidiii'j 

iif  disease  is  Miiiiilit   fur  iai'jelv    li\    the  eiiipiricaj  methiid  :  n, I\-.  Ii,\    the 

eiidea\n|'  tn  see  and  ex.imiiie   iiinii lalile  iiatiiiits,  tn  diaiiimse   the  ejise 

acciirdiiiL!  tl)  the  unnipiim  ,■(  the  si.jns  and  syniii'"iiis.  and  tn  treat   it  liy 

the  prescrilied  niethnds  nf  e\periciicc.     S m.||  |i;,s  tl)  lie  liaiiied  and  sn 

HMIch  has  In  lie  seen  dlliin'^  tl linieal  \eais,  that   the  student   uIm's  little 

ihniiL'lit  fn  the  natiii f  the  functi.iiial  distiirl),in<i'  wliii'h   is  respmiNihle 

fur  the  syiiil'lnliis:  he  fails  to  rcali/.'  that  after  all,  there  is  nn  esseii- 
lial  differenee  lietueen  the  eoiiditioii  liroiii;hl  ahoill  in  his  patient  liy 
Slime  patlinlo!;ie  lesion,  aiid  that  which  may  lie  produced  in  the  jahorn- 
lory   hy   experimeiit.il    procedures,   liy   dnnrs   or    1).\    toxins.      It    iiii!.,!    of 

course  he  reeoirni/ed  thai  Just   as  the  seien f  medicine  nrisrinated  h.v 

the  i.'rnupin<f  of  >\mptoms  into  more  or  less  cliaraeteristie  diseases  for 
which  the  most  favorahle  nntlind  of  t  re;it  ment  li.id  to  he  disen\ered  liy 
experience,  mi  nnist  a  ci'itain  p.-irt  nf  the  medical  traininj.'  he  iiiore  or 
less  empiriciil  hut  it  -.honhl  al  the  s,-iiiie  linic  he  rcili/ed  lliat  such  a 
iiiulhod  is  oiil\  a  means  to  an  end.  and  ihal  the  real  understanding  nf 
disease  can  he  ae.|uired  only  when  every  ahlioiiiial  eondifinn  is  inter- 
preted as  a  primary  or  secondary  conseipience  of  some  perv.'rted  ho.lily 
iMiictioii.  and  when  the  lr,iininu  in  .iliservation  and  the  inductive  method 
is  cai-rieil  from  the  hilior;itiii\    into  Ihi-  clinic. 


I'KI  I   \<  I 


•'     I-    1 ll>taMl     IXIHI  |(  ll.'i-    ,,f    rllliirlll     lll>tP  llcl.ii,     .y  li.l    \M>lll<l     l'lll|llli> 

••••iflllitir  IilIIiihU  III  lllstllli'lioh.  tliilJ  tll.y  lillil  the  >l  llili'llt-.  imt  o|||\ 
illllitTllrlll     li.    nil    lllllll\si>    1.1     lll.ir    Cil*.*    I'lolll    till'    I'lllK'iKIIIrl    ■.tillMlltnill 

liiil  jil>i(  that  tlii\  iin  too  iiiii(|i'i|iiiilrl\  pri'imrcil  in  t'iili<lniiiriitiil  |i|i.\«. 
ioliiuii-     l«l|n\\  Icdui',     |i>     liiilUi'     till'    (illillv^iH     |tossilp|i'         Till'     >llli|rMl     lllll> 

liiHr  ii  >ii|iiTlh'ial  ii.'.|ii taiM'i'  with  tlir  main  f'ailN  ul  |iliv>i..|"uii-  >ri.ii.-i 

hilt   liaM'   failiil  to  ai'ijiiiii'  ll ni|iiiriiitr  lialiif   ol'  niiinl   wlii.-h    will  I'li 

nlili'  liiiii,  thioiiuli  I. ■tin  lion,  i-oiii|,jirisoii,  ainl  |>ci>oiial  i  .•seairli,  to  iip 
jilv    till'  Knowlrijir,.   in   inai'tir'al   imiiii-inc  ami   siirmrv, 

I'of  this  lacl>  o,"  iM.ii.'lalion  lirtwcrn  itif  li;lioiafor\  ainl  rliiiical  stinl 
i<s,  fh.  i'lini<-,'il  iiiNiiiirtors  ail'  not  almif  rcopoiiHildiv  The  liilionitory 
roiii'sfs  III.'   i'i..|iii'ntl.\    iri\.'ii   without   ati.\    attempt  I.cintr  imnli-  to  >lio\\ 

Ihr  stmlmil    thi    l.caiiim   of  tlic   siilijcct    in    thr    inlrrprctati t"  iliMWIsr, 

Of  to  train  liiin  -o  that  in  hi,  hili-r  yrais  h..  may  \h-  aM.-  to  inlapf  l1i«' 
m.tho.ls  of  in\i  stiiiation  w  hi.li  ]„■  irariir.l  in  thr  lalioiat<ny,  to  the  stllilv 
oT  morln.l  ■■oinlition.  It  is  srlf  ,.\  i,|,.nt  that  ( \\  ithoiil  any  kiiou  hvltf.' 
ol'  .iisrasr     tl..'  .At.nt   to  \^llil'h  th.'  stn.li'iit   in  tlic  rafli.T  years  of  f)n- 

<-onrv  I'oiiM   Im'  .Apcctr.l   to  appiviiatf  III.'  rlinical  siu'iiilica f  wlia 

hf  I. 'arils  in  tlir  lahoiatorv    is  limit. •.!,  I.iit   this  sh.nil.l   not  .I.'ti'r  tlic  in 

stnntoi'    I'm. I, I    in, Ii,. at  in..'    \(  h.'ii.'M'r    I an.    tli.'    L'.'ii.'ial    appli.'atj.ni    ..f 

s.-i.nti(i.'   knowl.'.lyv   in   ih.'   int.rpi<tati..n   of  .liscas..!   .'.in.lilioiis.      r.iit 

III.'  .'hi.'f  i.'Iii.'.In    ,,f  th.'  .'vil  nn.h.iiM.'.ll.v    li.'s  partiv    in  tl nitiniiai 

ol'  .'irtain  of  ih.'  lain  r;.  .n-v    .•onrs.'s  int..  tl lini.'al   y.'ais,  ami   partl\ 

HI  th.'stn.jy  ,,(■  in.'.li.al  litiratiir.'  in  uiii.'h  th.'  applhatioii  ..f  i.'iysio|o._ry 
an. I  l.io.'h.'iiiistiN    in  the  pra.-ti.'.-  ..f  i li.-in,'  is  .inphasi/.'.l. 

V..l"ithstan.|in._'  ih,'  siifj.'i.nt  inimK.'r  ..l"  I'X.'.'ll.'tit  t.'\tl.o..ks  i  ,  plixs- 
lolo'-'y  availali!.'  ;.,  ili,.  nn'.ii.-al  stn.lrnt,  tliorc  is  iion.'  in  \(|iicli  jiartic 
niar  .iiiphasis  is  lai.j  upon  th.'  applii-ation  of  the  siiLj.'.-t   in  tin-  roiifiiic 

I"''""'"'"'   "'"   ' lii'iiH'.      In    tin'   jn-.'si'iit    \oliiiiii-   the   att<'m|it    is   miidc  to 

m.'.t  sn.'h  a  want.  I.v  iv\  i.'win./  Iln.sc  p.. i  lions  .if  p|i\  si.i|oir.\  ainl  liio- 
.■h.'inistr.x  w  hi.-li  .•\|M.|i.'n.'.'  has  shown  to  he  of  rspi.^ial  vain.'  to  th.' 
-•lini.-al    iiivi'sti[iat.,r       The    w.irk    is    not     int.'inl.'.l    t..    Ii.'    a    siil.stitntc, 

.'ilh.'r  r.ir  the  r.",'iilar  t.'\tl Us  in  pIiysio!oi;y.  or  for  tli..s.'  in  fnncti.inal 

|>ath..l.,.jy       It    is  siippi,.|ii.'i,tary  l.i  sii..))  v..lnm<'s.      It  .lo.'s  nol   start    lil<.' 

•'"•    ' '"■'"    '''^t    ill    I' ti.-nal    patliol..'.'y,    with    a    .■..iisi.l.'iati.ni    of    th.' 

' I '^' ■•'"•'■''  'i'i"ii.  iiii'l  lli''ii  pi'M I  to  aiialyz.'  th.'  possil.lc  cansi's  aii.l 

.•.iiis('.pi(.n..s  of  III,,  ilistiirl.an.-.s  ..f  fnii.-tion  wliich  this  fxhitiits:  hut 
it  deals  with  tin  pi .  s.nt-.lay  knowl.'.l'j.'  of  hnniaii  physiol.n.'y  in  so  far 
as  this  .'an  Im'  iis.'.l  in  a  •^.  n-ral  wav  to  a. Ivan.-.'  the  iin.lerstan.rm!.  ..f 
.li-^.'as.'  In  a  s.ns.  it  is  ih.'r.f.n  .•  an  a. Kan. -..I  1,'Xt  in  physi.>|o.jy  lor 
""'^'-   ■•''""!'    '"   'II'''''    ii|"'ii    lli''ii-   .'lini.-al    instill. -ti.iii,    an.l    al    th.'    sani.' 


I'tii  r\i  I 


VII 


liiiii',  II  n'\ir«  I'lir  lliiisc  <i|'  11  iiiiiliM'i'i'  fliiiiriil  i'X|Hiiciii'i'  uliii  iiiiiv  ih'siii' 
III  wrk  till'  |ili\  siiiliiuii-iil  iiili'i'|)i'i>latiiiM  of  iIIhi-iim'iI  <-iMii|itiiiti<< 

III  iiltini|>liiiu  »ii  I'lillil  tills.'  iTi|iiirfiiii'iit«,  it   iiii'«  Iktii  'I.  . m.'.l  r,>i.||. 
tiiil   to  (to  liHi'k   to  till'   I'liniliiinciitiils  of  Ihi'  siilijii't,  iinil  to  •'Xplaiti   ai 

xi'iply    lis    iHissililr    till'    iihxsicaj    aliij    plix  sicorliciiiirai    pi  iiicipli's    ii| 

wlii.'h  SCI  hnK..  n  pint  'if  plivsiolMi.'ii'al  know  li-ilac  tlfpciul'*.  I'liy.siolngy 
may  hi'  .•iiiisi.j.Tiij  as  ;.|i  applimtioti  of  tin-  known  Iiiwh  ami  farts  of 
phxsifs  ami  I'lirmislrx  to  I'NpJaiii  tin'  fiim-ijoiis  of  li\iiitf  iiiatti'f,  ami  it  is 
only  after  the  iMnit  to  u  liicli  this  application  fan  lir  iiuiili'  lias  lu'i-ii 
appit'ciatiil.  that  thi'  kiio\»  Ii'cIkc  may  lie  iisi'il  to  s>-v\r  as  tlii-  foiiiiilation 
upon   whl.'h  a  slip!  istniftiiir  of  i-lihiial   kiioulcluc  ran  In-  Itiiilt, 

III  onln-  that   thr  \oliiii juht    In'  niainlaiiii'il  of   irasoiiahlc  si/.',   it 

•i"''  l"''ii  ' issaiy  to  s.'l.-.-t   nitaiii  parts  of  tlir  siilij.'ct   for  partiriilar 

riMphasis,  the  liasis  of  si  h.-lion  luinif  tlii'  deifri'i'  to  w  hii-h  oiir  know  Icdiri' 
'li'arly  shows  ih..  valiir  of  tin'  applii'ation  ,if  physiojoujral  iiii'thoijs  lioth 
"I  oliscrvatioii  anil  of  thmiuht  in  tin'  stinly  of  ilisrasiil  romlitiinis.  This 
lias  III. I  lirni  ill, III'  to  till'  I'Xti'iit  of  oinitliiiu  lln'  appM'i'iitly  Irs.s  I'ssiMitial 

parts,  for  tli.'sc  havr  I n  tnati'ij  in  snlVn-icnt  ilrtail  to  link  thr  others 

loiiitlii'i'  so  as  to  preserve  a  loyieal  eom  innity,  ami  show  the  liearint;  of 
one  lielil  of  kliowleilire  on  another.  There  are  however  eeltain  parts 
of  the  seieiiee,  partieiilaily  the  physiology  of  iier\e  ami  niiisele,  of  the 
speeial  senses,  ami  of  reprotlnet ion,  for  whieh  applieatioii  in  the  ueneral 
liehls  of  nieilieiiie  ami  siii'jeiy  is  limit)  i|,  ami  these  parts  have  heeii 
oiiiitleil  eiitiivly  jt  has  I.eiii  jiiilm,!  |||„t  this  perhaps  somewhat  arl.i- 
Irary  seleelion  i,  jnstilieil  on  the  uroiin.l  that  the  or.liiuiry  text  in 
physiolo'jA    eoMis    these    snh.ieets   snniii"nt  l.\ ,    except    I'or   the   speeiiilist, 

lor  whom  on  the  othn    liaml ii'|ei|nate  review    wmi'-l  have  lieen  pos- 

silile  within  the  limils  of  sinh  a  xohiiiie  as  this.  With  reference  to  l.io- 
ehemislry,  no  altempt  is  nunle  to  review  the  pro])erties  or  ileserilie  the 
eharaeteristie  tests  of  th"  various  chemieal  iiiL'redients  of  the  hoily  lis- 
snes  ami  llniils.  This  is  already  siiftieientlv  done  in  the  texthooks  on 
Inoeheniisiiy.  and  in  the  niinieions  manuals  on  elinieal  methods.  I'.io- 
ehemieal  knowled-e  is  treated  rather  from  the  iihysiolosfist 's  stand- 
point, as  ai;  inte'jial  |tart  of  his  snh.jeet,  partieiilar  attention,  neverthe- 
less, hiiiii.  paid  to  the  far-reaehin.j  applications  of  this  latest  nejiart- 
menl  of  medical  seiemc,  in  the  elucidation  of  many  ohsciire  inolilems 
of  clinicil   medicine,  smli  as  those  :,f  diahctes,  nephritis,  acidosis,  uoiter 

and   myxciicma.     T akc  tlie   \olni f   \aliie   to  those   who   may   not 

!ia\c  had  time  or  opnortnnity  to  familiarize  themselves  with  the  techni- 

eal   methods  of  the  physiolo',iist   and    liiochemisf   as   used    in    tl d.rn 

elinie.  a  certain  amount  of  space  is  devoted  to  a  Inief  description  of  the 
methods  that   appear  at    present    <o   he   rccci\  inu-   most   attention,  and    to 

lie    nf    !;re;ltl'st    \;dnc. 


I'iiijilly,  it   >lii>iil<l  lie  iitiniicil  tliiit  tlic  |)iiiici|)lcs  of  sciiini  diimiiiisis 

iiiiil  lli('i;i|iy  :ilc  oiiiilti'il,  siinv  tlli'sc  liclnin;  In  ;i  li'iLilily  s|)fci;ili/c.|  sciciici' 
ti'i|uiiiii>;  nil  iiitciisixc  t  r.iiiiiiiv  "f  its  own. 

Til  tlic  lie;-;-  ,liMt  tlic  vciliiii,.'  iiiiiy  lie  iiisl  iiiiii(iil;il  in  ■■irmisiiiir  siiflicii'iit 
ilitci-i'st    t(p    stiiiuil;itc    M    iiMiic    iiili'iisi\c    study    (<(    tli.'    v.'iiiniis    suli.ji'cls 

Mliicli   it    iiitiixliiccs.   .1    liiii'f  liii(lii>i:i;i|i)iy    is  -i\,.|i   ;il    ll iid   of  c.-icli 

scctiiin.  T]ic  icffrciiccs  si'lci-tcd  iirc  tn  |i;i|icrs  tliiit  ;iii'  iiiciif  |inifi('- 
iilarly   known   to  tlic  aiitlioi-:  tlii'y  .-irc   imt    m cisNiirily   lliu  must    inipnr- 

tllllt     |ill'ilic;itiipr  i    (III    till"    sillijci-t.    lull     iiri'    ol'tiMI    clinscli     liccillisc    of    till' 

iisct'iil  r.'vicws  (if  incvioiis  \m.iU  tni 1  in  tliiMii,  i;itlicr  llijin  lii'ciiiisc 

"f  tlii'ir  own  <pri<:iii(ility,  Smiic  of  tli.^  piijii'is.  liow  .x  cr,  ;ii.-  icfiMTcd  to 
!is  iiiitlmrity  for  st;it«'iMi'iits  of  fjicl  wliirli  nuiy  .•iioiisi'  in  the  iciidiT  m 
di'siri"  to  jioiidor  for  liinisdf  tlic  ex  idciii-i'  iipon  ulii<'li  tlirsc  jiii-  IiiisimI. 
Till"  irfcrriK'cs  arc  usually  divided  into  two  !,'ioii|is.  "  iiionnt;r.i|)lis"  and 
"oii-rinal  jiapcrs,"  and  it   is  only  occasionally  that  specific   ivfcrciicc  is 

made     II  til,,   fornici'  in   tlie  context.     Tl iii.niial   iiaiicrs,  .11   tlic  otlier 

liand.  :\vv  referred  to  liy  .inniliers.  AVitli  tlie  <.'ciieral  field  of  tlie  siiliject 
so  ivell  covered  Iiy  siicli  excellent  textliooks  as  Rayliss'  "Principles  of 
t.ciieial  I'liysi.iloiiy,"  St, •wait's,  Tlou  ell's,  Starliiiu 's,  .ind  Ifaliihiirton  "s 
••llniiian  IMivsioloMics."  and  Leonard  Tlill's  "IJeccnt  and  T'lirtli.'r  Ad- 
xaiiccs  ill  Tliysiolo-y,"  tlic  author  has  felt  free  to  |.icl<  aiel  hooso  from 
the   iiioiioi.rai>Iis  and   original   i)apeis,  topics  that    arc  ordinarily  iiassod 

over  cursorily  in  the  texthook.  and  when  this  has  h 1  done,  the  refcr- 

•■' -^   ii''<'   siHiicwhat    re   extensive.      Siudi    is   the    casr.    for   example    im 

the  ehaiitcrs  r.datiiii;  to  t! heiiiistry  of  i-.'spirjitioii.  to  the  mclal)olis:ii 

of  carhohydiatcs  and  fats,  to  the  jirohli'iiis  of  ilictcti,-s  and  liiou  th,  to  the 
physiiM.cheiiiical  hasis  of  neutrality  riiiiilatioii  in  the  animal  hody,  and  to 
I  he  action  of  enzymes. 

.\ckiiowlednr„ieiit    is    u'latcfully    made    for   the    assistance    ami    advice 

"I   ''"■  pieparalii f  the  hook,  part i.-iilarly  to  Doctor  1{.  (I.  I'earco,   for 

'' oiitiiliiitioii  of  .several  chapters,  to  whieli  his  iiaiiie  is  attaidied.  and 

for  which  he  is  entirely  respoiisilile  ;  and  to  Doctor  E.  P.  Carter,  m  hose 
eriticisiiis,  after  patient  perusal  of  the  iiufinishcd  niauuscript,  -vverc  of 
inestiuiahlc  valiii'  in  its  final  revision,  .\cknow  Iclirnicnt  is  also  uiarle 
to  Doctor  |{.  W.  S,-ott  a, id  l'roless,ii  |-'.  K,  |,l,,yd,  for  valiiahle  criiieism 
and  advice,  and  to  the  former  for  a  chapter  on  the  "Clinical  .\p|>lica 
tion  of  Klectidcardioeraphs  ••  To  ^[iss  .\ehsa  Parker,  ^T.\  .  the  author 
owes  a  '_'re;it  delit  of  <rratitiide  for  thi'  thorou'.'h  and  i)ainstakiii<,'  w;iv  in 
whi.'h  she  prepared  the  niauuscript  f<ir  the  press,  and  for  her  never- 
tirinv;  endeavors  to  ha\c  the  spoUiii..'  and  imnefuation  in  conrorniity 
xMth  Wchstcr's  Di.'lionarv.  I'or  assivl.mee  in  the  |iieparation  of  the 
index    thanks    are   due    \<,    Mi-s    .M,-iriaii    .Vrmmir    1    .Mrs.    .Xfaid-'arlaiH. 


riii:i-\(i: 


IX 


iiiiii  for  ppriiii.ssioii  lo  use  ccrLiin  of  tlic  li^'iiit's  iiii<[  illuNtriitioiis,  to  the 
various  autliors  and  pulilislicrs  who  "ir'aiitcd  it.  l-'or  llic  cxccllcMt  iiiaii- 
asiiMiiciit  and  carcfiii  cxcculion  oi"  llie  picsswork,  the  aiitlior  «  isiics  to 
thank  tlio  ]>iililislicrs,  whose  cointcoiis  ami  i'ricinlly  dealings  liavc  alwa\s 
iriadc  thr  wurU  easier. 

.1.  J.  I{.  M.u  i.i:ni.. 


riiivrrsity  (•!'  Tcionto, 
Tiinnitd,   (ViiKtiia. 


CONTENTS 

I'AKT  I 

TMl-:  I'llVSK  OCIIK.MICAL  itASIS  Ol'  I'll  VSK  »|.()(;i(  AL 
I'KOCKSSKS 

('IIA1TI:K    I  ,.^,,^. 

•  iKNKUAI.     (■iin.siiiki:atiiins , 

The  I,ii«s  (if  Snliiticii.  :;:  (i;,s  r.;i«.s.  :;:  ()sra„ii,.  I'],>«iiic.   I:  I'.iul,,;;!,;,!  >r.'tli".ls 
fur  Mc-i^^iiriii-  Osiii.,ti,-   I'lv-snrr,  li ;    llfiiii.ksis.  7:    I'hiMii.il.v^is.   *. 

CIIAI'TKIt    II 

Os^rnTlr    l'i;i:ssii;h;    (C(p.\t'ii ^(^ 

.Mc-isiiriMiMMit    l.v    I)c|,rr>si..ii   ..f    Kiv./.iim'   Point,    lu;    Tl.r   Hdr  of  Osiridsis    Dit'- 
lu^inii.   ;,i„l    AlliiMl    I' essrs    111    I  Ml  \>i(,l(.-ic    MriliMnisiiis.    11. 

CHM'TIK    III 

Kl.KlTHll      C'lMirrilviTV:     1  )lj~SO(l.\T|nN ,      \\J,     I,i\|/.\TII)\ IC 

lii<l|ii^ir;il    AppliiMtiiills.     111. 

•  IIAl'TKK    I\ 

Tin:  ri:iMii'i.i  s  iNvoi.MiMN  TiiK  I)i:h:i;minati()n  ut   1Iviii:(|(,i;n.i,is-  Cum  i.siii.viioN   JJ 
Titr.-il.lo   Ai'i.lity   iiii,l    Alkalinilv.   -J^:    Ailir.il    Dryirc   ut'    A.i.litv    iir   Alkiiliiiitv. 
il:!:   Miiss  Actinn.  ;;:;;    Ai.plii'iitiiiii  t"  tin'  .Mi'.-i.sinriin'iii   .pT    Iliun  C.in.iMitratidii. 
L'll;   Aiiplii-iitidii   in   D.'l.Tniiniii;,'  tli.'   !i.;il  SHvn;;tli   ,.f  .\.-i.ls  or   Alk.'ili.s,  lN. 

CIIAI'TKR  V 

TlIK    rill.VClCI.lS    INVOI.VKII    IN     IIIK    MkasI  i;  lAl  KNT    nf    IImm<.k,1;\    hiN    ( '|)N(  KNTKAriON 

I  <  "on  T  "ill .1,, 

riif   Klivlii.'  Mo'lin.l.  :;!•;   Tlir  Ilnli.-Mtcr  .Mctllil.l.  .■■•J. 

I  IIAl'TKK  VI 

Kli.riATln.N     ,|K     .\KlTI,Al.ri\       Is      nil.     A.SIMM,     |1..|,\        \M,     A.(  Ilinsl.S -.X 

r.uflVr  Sul,s!;.h.-,.s.  ■.\C,-  Tlu'niv  „f  A.'i.li.sis.  :> :  M.Msiircni.'nl  uf  ll„  !!,.>,. u,- 
Alkiilinily.  M  :  Titinlimi  MwIuhIs,  tl:  Co,  cMinhiniii-  P.uv.t.  Il';  !ii.lii,-.'t 
Mi'flioils,  -lii. 

CIIAI'TKH   VII 

CoT.r.OlDS 

ni.-irru-t.ri.ri.     I'r,.j.,.,  li,-..     ,-„, ;     CimnMl,.,  iM  „s     „r     Tin,.     Cui'loiilnl'    S.,luti.'.„.!    ' 
">1;    Tyn.hill    I'licnnniom.ii,    .-|1  :    KoliiliM'    In.litri.Mhility.    ."il  :     i:ir,i  ri,.     l'r.,|„T- 
til's.  .-,.-,;    ItiovMiinn    M.iM.niiMil.  .-.7:    Osnn.ti,-    I'lvssiirc.   .-|7. 

xi 


■iM  C'ONII  SIS 

rilAl'TI.K    \|II  ,.A 

'hi  l.olJiH     (('iNl'ji' 

Sus|...iis(,i,l,   ;,ji.|    KiMiilsui.ls.  (Ill;    (icl;itiniA-iti Ill;    lnil.il.ili..ii,   CJ;    Ac-liiiM    <■( 

i:ii'i'ti(ilvliM    .,11    Cilli.i.ls.    li::;     I'r.itiMUs    !is   Cillniils.    (i:!;    Siirl'iicc    Ti'iisi.ili.    lil; 
.\ilsor|.ii,iii,  ri.">;    l-.xoix.liiv   Ki'iicliiiii.s   Di  | liii;,'  mi   Ailsiirjiliiiii.  tl(i;   Oin.litioiis 

!llrtn.'M.-il!-     ..r     Illl!ui'llrnl     l,_v     A.IsilI  |.t!!l!!,    I'lT;     I'liysi,,!.!-!;'     1'i-:ih;--;h'S     Dl'l'Cll^lil!^' 
Mil     A'l-Hl|'lioll.    (i'.i. 


t'llAl'Ti;!!    I.\ 


I'l  l;\1 1  \  I  s.    Ill;     \'.s/\  Ml  : 


Till'    Niitiii.'   I'f    y.h/\ Ac'tinn,    7-;   I'mpcrtics  uf   Knzvim's,   7:1;    Ki'viTsiliility 

III"  Kii/yiiH'  All  ion.  77;  S|ioril'n'ity  of  Kiizyiiu-  Actinii,  7!';  ririiliiiritii's  ..f 
llii/yiiii's.  Ml;  Tvpi".  .<(  ]',t\/.\mo.  M;  ?:ii/_vnii.'  rii'|inratiiiiis.  '^1';  ('oiiiliti"iis  f.>r 
l!ii/Miiii-    Aitivilv.   S;; 


PART  II 
TIIH  (IKM  ri,.\Ti.\(i   I'MIDS 

CIIAITKU   X 

111, ■Kin:    Irs  Cinki.m,    I'Knri  i;tii:s    (I'.v   1{.   (1.   l'i..\Kii:i S.'i 

(^'iiiiitily   .if    I'.l 1    ill    till'   lioiiy,   s:>:    Wiitcr   ('..utriit.   Sil ;    I'mtcius.    ^7 ;    1-\t- 

IIKIlls    lilhl     Alllil'rllllc'llts.    S!». 

ClIAl'TKK    XI 

'rm:  I'l.iioii  Chi.s   i  I'.v   '{.  (i.  I'l  vki.:) <il 

Ki''l   I'.l I  ('i.il'iisilcs,  1.1    KiUl Ui's.  ill;  ()rij;iii,  iij  ;   Hati-s  1.1'  |{4';;i'iirratliin, 

!i:::   llciiM.ly^is.  I>,"i;  I.,'ii.-iH-\ti'>i.  '.M'l;  l!l 1  I'lalclcts, ',I7. 

ClIAl'TKK   XII 

Hi, Hill.   I'l.. I  MM, IIS 

\  imI.I.'    rli.iii-.'^    ill    the    l;|<M..|    Diiriim'    r|..tli!i^.    liS;     M..tli.uls    i.f    Kctaiiliii;; 

('li.liiii-.  '.i;i;   N;il f  tin'  ('•|..ttiii;i  I'lmos.  101;   luthioiico  of  Calcium  Salts. 

in:'.:    liilliiiiiii-  ..t'  TissiM'S.    101. 

•  'IIAI'TKI!  XTII 

r.i....ii  ri.i.iiiM,    -c.im'i.i ]n,: 

■rll.'..|i.>   I.I'    l'.|."..|   ('[..llill;;.     il'ii;    Ililr;i\;,>.ii|.u    ( 'I..!  I  ii,^',    lll7;    MwiMllClllclll    '>( 

III.' I'l. .Hill;;  Tiuii'.  Ins;  r.|..,.,l  ('lotting  ill  \'ailiiiis  riiysii.li.nic  OiiiJlli..ii.s,  llii; 
r,l..<i.|  ('L.triiij;  ill   |)i>.;isc.   Ill;  III  iiimrliayii'.  Di.soasr.^',  UJ;   Tluoiiil.us  Fuiiiia- 

tinll,   11.1.  .11 

•IIAPTKK   \!V 

I.V\fl'jl      r..|:\l  \l  li.S       A.Ml      (,'11,111,  \  I  111- ||-, 

(m'ii.i;iI    r..ii<i.|.,|;ili..ii<.    I  1 .-, ;    i:s|.,,riiii,  iiliil    I  inr>l  iuiil  i.  iis.    11'^;    IMi'iiia.    |J(i. 


ION  IIAIS  Mil 

I'AKT   III 
<  llfCI  j.ATIoN  (»|.-  'IIIK  l!L<M)|» 

t'UArTKK    W  pM.r. 

r.i.O'M)    ri:Kssi  i:i: VJ-J 

Till'   Mi:in    Ailni;,!    Ill I    I'lr^siin-.    Ij:: ;    Mm-iiiy    Miiiinnii  In    Ti  :i.in--,    IJ:;: 

S|.rili:;    M:iMcinii'li'l    'I'lu.  ili^s.    iL'l);    Clinirill    Mricsi r:rlll-.    IJ^. 

r||\|'Ti:if  Wl 

'rill.    r.M   |o|:s  (•ii.S(  lUM  h    r\    M.MSIMMNC     Till     Ulooii    rKKSSI'lil W'.l 

l'lllii|.ili;;    Ai-liiMi    "f    tlir    lliilll.     l.':l;     I'l  ri  |'liiT:lI    Kr^istiilhi'.     I .;  I  :     Al it    ul 

l!l I  III  the  ll"il,v.   l:;."):    KlVi'.ls  df   lli'iiiMiiliii;;!'  :ii;i|  Triiiisfiisiun.   I.;i):    \i>i,(i<- 

il.v    ..f    till.    r.|..(i,|,    llll;     Kill  stir  it)     i.f    Vr><rl    Willis,    111'. 

(■ii\i'Ti;i;  .wii 

Till    Al  vi.i\   (||.    Tin;   llKMir lit 

Till'  riiiiipiii:.;  \rtirin  nf  till'  llciiil.  Ill:  liiliiii  ii.jiii.-  I'li'ssiiiv  CiiiM'^.  III!; 
<'oiii|'iii  ivun   III'  llll'  Cmim-  .    lis. 

•  ilAI'TKIi    -Will 

'I'lii     I'l  mimm;    Airi"\-    ni     nn.    Ukaih     i('i(\i' |."il 

•  ■iiiitimr  (if  till'  Intrai'iiriliiil  I'M'smih'  Cimm's.  l.'il;  \  fnt  liiiilin-  Cuim'.  151; 
Aiiiiiiiliu  Ciiivi'.  l."),'!;  The  Xl'.'.liiiiiisin  nt'  Oiiciiiiij;  lunl  Closiii;;  ,,(  tin'  ViiKrs, 
l.")l;    Tlio    llt'iiit    Slum, Is,    l.'jT:    Cjiusi's    of    Siiiiii'ls,    l.")7;    Koi-inls    I'l'    Siiiiii'ls 

'    i;i.'itin|.|in||.io,;,|l|>,,     1.-^. 

CIIAI'TKU    XIX 

'I'ln:    NiiKiTiiiN    iiK     iiii:    lliiAur 1i!] 

\'\ I  J^'il'l'ly,   !i;i:   I'liliisiui,  ,,f  til.'  lli'iiit  Out.sidi'  tliL'  r.i.'lv,  Mi:   Ki'siisi'itii- 

ti.m  .il  till'  lli'Mit  ill  situ.  |i;i ;  Ki  liitinn-iliii.  iit  tlir  ( 'lii'iiiiiiil  ( •i.iii|..i-iti..n  nt  tin- 
I'l'ifiisiuii   riiii.l   ill  Culil  Mnoili'il  iinil   Will  lit  liliin.li',1  lloiirts.    liiri. 

fllAl'TKli    \.\ 

I'm  sioi  iM.v   111      iiii;    lit.  Mini  At I711 

«)ri';iii  iiihl  l"iii|.:iL;;iti..ii  nt'  tlii'  lioat,  170;  Mvo^niii'  nv[.i.tli,'sis.  171;  Xciiio 
HOiiic    llvputlicsis,   17J:    Till-    l'iii-.Mii!ikor   nf   tli.'    llnirt    iiml    Ili'jiit  I.liii'k.    171: 

I'liysi.'lu^ir  rii;ii:i.  I.'iisti.  -  .it  I'lii'liii,'   Mils,. I,'.   17i;. 

I'llAl'TKIi    XXI 

I'm  >H)I.1H,\     111      IIM      lllMMIllAi     ,r,p\l'l.i ,     )S2 

('ii;;iii  .-iii'l    l'riii.:it;iili,,ii   i.t   llii'   ll.-iit    in   tlir   .Miiiiiiiiiiliini    lli'aii.    1'>J;  lii,l- 

iii^   Tissiii-   ill   til,-   Mil iiliiin    lli'int,    I^U:    Sili'   ,.1'  ()iii;iii    ,.|'    |'..iil.    Is,. 

rilAl'TKK  .\.\1I 

I'm  sioi  ()i,v  1)1    nn.  lliMiiurAr  i('.i\t"iii 1:,1 

\|...!i'  ,.1-  I'l'ij.ii^iiti,.!!  ,.r  !!■.-  1^-it  in  Ihr  .\i;rii!,-s  :r.;.\  {:,,,■.,  [):r  .\;i:i,iis  t,i  Tin- 
Xriitri.'li's.  KM  :  SpiPii,!  ,.l'  r„'i,t  in  ili,.  Vonlriili'.  I!i::;  Kil.i  illiiti,.ii  ..(  {],<■  \'.ii 
liii-li";    1111.1    .\iirii'Ii's,    lii,". 


XIV  (li\l|\|S 

III  M'Tlli    Will  ,.^,;|, 

Till      Itl.nnlil'lOU      l\      I  III.     .\l:ill:ns ^  l<i^ 

Thr   I'lijsi-i.    l;i-:   Cm, .ml  Clinrii.M.  ri,-ti.  s,    I'.iv;    1{,-,|.'  i.f  •ri;iii>iMi-^i (    I'liK.' 

WiiM-.    I'.'s;   (•..111.. in    i.r  till'    I'liKr   Cuiv,-,  jiiH:    \,|..,.it\    |'ii|>,..  I'liH;    I'iil|,nl,|,. 

I'iiIm'.  I'm':  Aii.-ilvsi.-,  ..f  til,.  Curv,',  i;ii:;;  Til,.  Di.n.ii,.  \V;iv,..  :;ii:;;  ('..iii-..^  .,f 

l'i.':i|.|.,;ii;iti.','  ..!'  111..   I'nU..  in   il,..   \',.in-i,  Uii.". 
••11  M'l'l  |{    .\.\l\ 

l(\TI     ..|     M.iVrMI  M    ,.|.     i  hi:    |',|.i,i|,    |\    y,,,     liiDMIi    \i,-sn.s JlliJ 

V,.I...-it,v  ..(■   |-|..v,,  I'lKl:   M.,--  M.,u.ii,.  lit  ..('  tl...  Ill I.  Jux;  Tli,.  \"i.-.-,iiil   Hlim.l 

""»    i'l   ^' -I-':    \V..ik  .,r  111,.   Il,-i,,t,  ■Jll';   (•ir.iil:,ti,.ii  Tim...  L'K::    .\I,iv,iii,.tit 

"1'    111...,.!    in    111..    \-.  iiiv.    L'l  I. 

I'll  M'TKIf   .\.\\ 

Till      ('ONIl:,)l.     ,,'       Till.     I'll.-iTI    Mh,\ o|,; 

.\,.|\,.   (',,i,t...l,    -17:    \:,-ii<   (•..iiin.l    in    tli.^  C.l.l  1.|,m,.I,.,1    i,m,|    Hi,.    MiuiiiMiilinii 
Heart.  i:i::  T..iii.-  \:ii;iis  .\.ti..ii.  l.'-_M;    .MlVn.iil    Vai;iis  liiipiilsi.s,  ^.'L':.' :    M,..  Iian 
ism  ..I'  \:i;;iis.  :.'!' |  ;   T,.|iiiiii:ili..ii  ..I'  ih,.  V.,-ii<   KiU.i-  in   tli..   ll..;irt.  L'L'.'. ;   Svm 
I'litlioti,'  ('..iili.il,  2-27. 

ill.M'TKi!    .\.\\  I 

Til.    r,)\  I  mil.  ,.K   1 1  I    I'll;.  1  1  M  |..s    ,i'..M.'iii ^j,i 

\,.n..   r,,iiir..l   ..)■    r,ii|i|iri:il    l{,.sisl.iii,.o.   L':.'li;    Di't,'.!  i..ii   ..t'    V;is (..r    l-'il.,.i> 

111    N..M..S,    L'.;i  :    ()ii:;ln    i,(    \;,v,,,,i.,t,.i     ^■,l^,     Kiln. is.    2:;^' :    >'..i- t..i     N,.m.. 

I',. lit,!-;,  L':;.'i;    1ml, p,. II. I,. lit   Tmii.ity  ..T   l!l.i,.,l   \-,.><,.|s.  i;.':il. 

I'll  \i'i'|.;i{  .wvii 

Tiir  I ',is  ri;oi    ..t    iiii.  I'li:,  i  i,\  1 1..\    .('..ni'mi j..',7 

('..nil.. I  .iT  tl...  \';is..m..t..i  (•.■iit..r.  2:'.7 :  Miinniin,'  ('..n!i..l.  :;::7  ;  NVu,.  (•..nti.il, 
-■:-■:  I'l.-s.ii  1.11,1  |)..|.i...<s..|.  Im|,iiN,.s  -:'.'.!:  I{....i|.rii,.;il  liiii,.iMiii,.n  ..f  X'ns.iil.-u- 
\',..i-.  -Jl:;.,    Iiiilii,.|i..,.  ..f  (limitv  ..n    ili,.  ('ii,iil:,ii,.ii.  -Jll. 

I'll  \i'Ti;i{  .\.\\  III 

l'l  ,II.I  M;nn  s    ,.!      r,l..i..|.     SlI-riA      l.-.     flllTMN     \'|S,1I!\ o|; 

rii.iil;,ti..ii  in  ih,.  liiain.  L't7:  .\niil..mi.-;,l  I'li-iiliaritii-.s.  :;ir:  I'li-sii'iil  Cin.li- 
li'.ii-^  ..r  rii.iiliiti..ii.  JlH:  \-,,-..iii,.!..,  N,.,v..s,  J.-,L':  Intra, ■iniiinl  rii'sMir,'.  L'.'i.'l; 
<'ii.-ul:ili..|i  llir.i.|-!i  til..  I.!in-s,  L'.",:; :  ( •ii.-iil:iti..ii  tlii..ii:;li  Ih,'  l.iv.r.  -J."..':  Tin- 
i'..i..|iaiv    I'ir.  iil:.ti..ii.   L'."i7. 

i'ii.\i'Ti;i(  \\i.\ 

I'li.M.M.     Nri-i  II  MT,.N-    ..1     CiiirMN     I'm  .sKii.ixii,     .Mr-nil. 1I-, ^'.-,ii 

i;ii.  tr....ii.li..;.i;ii,is,   :.'.-:i;    Til,.    \',.i,tii..iil:.i.   <',,iii|,l,-\,   •_•(•,;;, 

I'll  M'TKIf   .\.\.\ 

l'l  I"  I'    M      Al'I'l  I'    M  1"N-    111     I'i  111   MS     KlIN  Sli.l.lli.l.      .\Il.TII..|.S     ,  l'.l\  I   '1.  I J,;,., 

HI, ..Ir.ii.iir.ii.. plains  ..f  ili..  .\I..i,.  K.>.ii;il  K.irni.-  .,(  Canliae  1 1  i(-iilariti,.s,  'M;: 
Sini:^  \rv!ivt!im;-,,  -ilil ;  s,-.,,  ! !  i  irli  rai  ilia,  l^'itl ;  Til,'  Kxtiasv,!.,!,  ,  ^iW, ,  l'ai„\ 
vsiiial  T'liliM-anll.,,  I'liU :  \i.ii.iil:ii  Kil.i  ill:iti..ii.  L'i!!i ;  Anrii-nliir  Kliil|..|.  Jii'.i; 
ll.ait  l.ld.'k,  :.'7n. 


COS'll  SIS  XV 

CIIAI'I'Klf    \\\l  l-Ai.t- 

C  'I  I  sic     ,1      AlTI  Ic    \  1  ln\^    ul     <  'l  l;l   M\     I'l  I  \  s|i  II  (H.  Il       Mil  lluns     ,  I  ■|1\  I     H JTil 

|'iil\  s|ili,\^iiiir;;r;iiii-i,  L'7-': :    NrMciiiM    I'lil-i'   'I'l.-ii  iii^-^,   L'7.;:    Siiiiiilt:iiii''iii<    AitiTlril 
l'iil~r  Tiiii'mi;^,  ■2:1;;    vKiioiiiiiil   I'lils,.,.  L'Ti'.. 

CIIAI'I'KK    X.WII 

I'l.iMiM,   Ai'i'i  ic  \  i  |i'\^  111    CitiMiN    l"ii\ -1111  iii.ir   Miiimiis   '('nsi'ni Us] 

Mril> llirlil     Ml     llir     \Liss     M.u  t-llli'lll     n(    ||i,'     111 1.    :.'>|  ;     Tl,,.     NuiIMIll     l'"lo» . 

jsj;  fiiir'.;,!  coniitini,-  wiiicii  Anv.i  ti„.  i!i iii.,u,  L'-::. 

(IIAI'TKi;    WMIl 

SHDI   K -J^J 

(li:i\it.\      SllMili.     L'S?:      lliMllirlllia;;,.    Miii.U,     :.'>^  ;      AlM^llirtir     SliM.I>.     :.'«s;     Spiiiiil 

Slji.rk.    LNs;     .\.u,,us    ^Ii.mK,    l'^;i;     Siii;;i.;il     Slin.-li,    I'Mi;     i;xi..'iiiiii'ntiil     In 

M'stiuiitioM    i.r    Sii.i.k,    L'*'.!;    Tic.-ilinciit.   ■.';i.'i:    f'liiiM'    Ml'    S iiil;n;v    S\  iiiptMins, 

-i'.'i. 

I'AK'T  IV 

i;i;si'ii;.\Ti(".\ 
ciiArrKU  .\.\.\i\ 

III  ^I'lKA  1  ION .,,,,, 

Till'    Mrclliilli.-    Ml     l!r>|.il,i|lM,|.    J!i:i;     |-|..„uii'    Mil. I     A Mil    .  1 1'    Air    III    lllf    l.:ill;iS. 

-'!i|.:     l{,.s|.ir;,t.irv    TiM.-iiiu-.    .■.n;:    Tip.-    I  iitr;i|.lriii;il     l'l.■^sur.■.    :;iil;     |  iitlii.'iii-.' 
'Ml  lil I  IMcsiir.'.  :iiii;. 

('iiAi'Ti:i;  WW 

Tin:  Mivii  AM. -,  ,11    I!i,,ni.M|,,\    ,(,im-|,,    ;|;\    ];.  ,;     |.,  >,,,  ,         .,,1, 

V:ni;iti..ii,   in    li,..-,.l  S|.|m-,..   I!,  >i,|ii:il   Aii    ii.i.l    ilir    \|i,|     ..,,,.1    \it:il   <•:,,,;,, iiirs   in 
\Mii..n-   I'll!  .i  i|M:;i.-  iiii.l    l':itli.il..:;i.-  I '..li.lii  |,,iis.   :;lii. 

•  11  \l'll.l{   WW  I 

Till.  MiiiiAVir.N  (II-    Kisrii;  M  io\    .  Cns  ;  'n  i    ,  [:\    );    (,     |>;  ^m  ;  .  o,- 

Tli,.   .\I.M-I,:i,iis,ii   ,.r   III,.  CIiiiml;.-    in   r:i|,:„-iiv    mI    i  It   Tl|..nu   :in,l    l.>,ii:;s.    :;i.->: 

Tin-  M„v,ii„„is  ..I-  tlir  Itilw,  :;|-:  T!i..    \,-ii i   ,|„.  M„s,-„|i,t,y,.  ,„•  f|„.  lin.s, 

-■:i:';  Tlir  A-ii.,n  .,r  ili,.  |)m|.Iit:ui,i,  :;-Jn  ;  Tl,.-   lUh-.u  ..f  tli,.  l{.-.|,ii„t.„-v  M,,,,- 
liii'iiN  MI,  til.-  I.iiiil;-,  .:i'.-,. 

<'ll  \l'|-i:i!    WW  II 

'I'lii     ••■•mi:"!.    .11     Iti  .^i'ii;MI-iN 

Th,.  i{,.s,.i,MT..,-v  Cpt.-i,,   ■■::■    u,,!,.,  .■„„ti..l  .,!■  tl„.   |;,.|,i,,,t.,iy  c-iit.-i,  :;:;i. 

CIIAITKK   .WW  111 

Till.     (•..sin.. I,     ol       li!  <i'lli  M  |..\       ilVisi'i,, 

•^'''''''"'  " '.  ■"•'■":  T.-iisi.iii  ,  r  .X).  .-111. I  ()    i„  Ah...,h,,  Air,  :i:;:i;  TrnM..ii  .,|- 

f'O    il,  \-,.|„,„s  |:i,„„i.  .-.i:;. 


\vi  (ii\ii\rs 

I'll  A  I' 1 1  :i!  \.\,\i.\  |.,wi> 

I'lu;  ('ii.NiKiii,  iPK  l{Ksi'irATii'\   ,('i>m'iii    ( |lv   K.  (J,   l>i;Anci:i .Ml 

IMiiiinIi f    till'    Alvi'Mliir    tiiiws.    ;;ll:    Mi'llnnj    Cur    Niiini:il    SnI.jrcM^.    ::i.'>; 

t'|ii,i,„|    M,.||,r„l.   :.\7. 

(•||\l'Ti:U  M. 

Till     iv.Miidi.   ()i     l!i  sriiiMi'^N    iC'iNi'iii      .     .  nm 

'I'lic    Niiliiii'   iif    the    Hisj.ii.itiiiv     II II.',   :;i!i;    Kchil  ii.ii«lii|.    LiMu.-ni    CO,   ..f 

liis|.iii><l  Ail  ■•iiiil  I'lilninniiry  Ni'iitih.l  Imii,  .'l.'ill;  I'uMsiMlily  lliiil  ri )  S|MTilii'ii||i 
Stiniuliilis  ill.-  ('.■iil.T.  ."..'iJ:  li.'hitii.ii>lii|.  iimihii^  A.i.li.>.i>,  AU.'.'hii  ("(),  ;iii.| 
l(.'S|iiriit..i.v    A.lixit.v.   '.''A, 

<  IIAI'TKK    M.I 

'I'lii    Cis'iKiii.  (11    Kisi'iKvn.iN    .••.i\i'i :',.",(! 

Til.'  ('iiii>taii.v  III'  III.'  .\l\i'<iliii  CO^  'ri'iisi.iii  iiMiii-r  .Nuiiniil  ('.iii.litiiiiis,  :J."iri; 
N'lisilivily  i.f  llir  Cititir  In  Cliiin-.w  in  tli.'  <  "i  >  T.iisi..n  nl'  tli.'  Alvinlar  Air. 
'■'''•'■,  .Mvi'iilni  (■<).  Ti'iisiiiii  iliiiiii;:  r>ti':.tliiiii;  in  :i  I'miliiiril  S|iiii'i',  ;!."i7,  in 
Uiiii'lii.l   Ail.  .".liii.  mill  ill  Aini.'ii.  :iill'. 

<'IIAl'Ti;i!    .M.I  I 

'I'lll      CiiSIKiil,    111      Kl  SI'lli  Ml.iV     lI'iiNl'lii :Mi 

Till'  i-:ir.it  iiC  .\iii~iiihii  i:\.'i.  i-.'  mi  tii.'  i;.-|.ii:iii.iii.  :;."ii;. 

I'llAITKI!    .M.lll 

Till     CiPNiiiiM.   .11     l!i . --I'll;  Mills    (Cist'iii .■:71 

ri'iii.illi'  liiralliiiii;.  .mI  :  Tvprs  uf  |',.|in.lii-  liiiiilliiii^.  :;7!  ;  I'aii^i'^.ir  riTin.li.- 
r.ii'atliiii;.',  :i7l'. 

i-IIAI'TKi;   .M.IV 

i;i  ^niiAiiiiN    iiM.iMi    riii:    I.im.s :;7s 

Tiaiis|'i'rl.iti.iii  111'  (;aM'>  !.>  th.'  I!I.mi,I.  ,';7!i  ;  Tiaii^innlal  imi  "(  <1\y^in,  .';7:>; 
|ii>sii.'iati.iii  Ciiiv',  .".^'l:  Dill'iTiMii'i'  lu'twi'.ii  Ciirvi's  nl'  Itliiiiil  an-l  lliMiinululiin 
Snliitiiiii.  ;>.".:    Kati'  III'   |)is<.ii-i:itiiiii.  .'ImI:    |)i^slll•iati..Il  <'iiiisiaii!.  :>s, 

IIIAI-TI.U   .M.V 

III  r.lMl:  M  111\     UIMiNI.     llll      I.IM,^     i('.i\l'lii .'IllO 

Mraii-    111    Wlii.'h   Ihi'    r.lim.l    C.iiiii's    llir   Ca^.'s.    i'.'.ni;    (l\.\uiii    |{.i|iiiir nt    ul' 

llir   Tissii.'^.  .';ii:;;    .M.-.-liani-iii   l.y   Wlii.li   thr    MiMiian.!-.  ..I'   llir   'l'iv<ii.'s    i.n    n^,y 

-I'll  All'  .\Ii'i.  :;ii7. 

<IIAI'I1;K'    .M.\  I 

Till.   I'm  >iii|  i«,\    111'  r.KiMiiiM.   IN  Ci.Mi'i:!  SSI  11  All:    wn  i\    li\i:rnili   An:      .      .     .   .Wi'.i 
.M.Miiitain    Si.kht'ss,    I'.li'.i :    ( 'iniijiri^M'.l    Air    .'^irkiii'ss     (('jn^sun     Pi'iiiiM' i.     It'L': 
i'rartii-al    A  |i|ili.ati"ii    in   Tii'atiii.'Mt.    Iml. 

I'llAITKIt    .M.VII 

'I'm.  I  "11:1  II.  Miii:\    .\-,ii   |{i.si'li;\  iiii:\    ('iiAsi.rs   .\.  iiiM  p  s  s  s  iNi;    Mr.s.ii.'.i:    i:\iiaisi     tin 
.Mi'i'lialiii-al    l':i,|,.i.    Iln;    .\.m,,'1~    l';„t.,i,    IIJ;    II. ni I'ailiir.    11:;. 


roNI'l  \TS  XVII 

r.M.'T   V 

i>h;i;sTinN 

fllAI'TKH    M.\lll  iM.t 

(JhsiKM.   rinsimiM.N    "I    uii     iMcjsrni    (ii.wns II* 

.\lirrM»|.u|pii-  <'li;iiim"<  .liiriiio   A>ti\ilv,   lis;   MiMliiiiiisiii  of  Sr.i.'tinii,    liici;  UthiM 

CmHioI.    fS\. 

CIIAI'Ti:!!   WAS 
riivsnM.iN.v  (u   'iiiK   liiiasTivi:  (ii.wns   (Cunt 'in tJ" 

lliiiniHii,'    ('iHiImfI,     I-."i;     Ni'IMHIm    <'iilitl.il    n!'    llii>    l'iillrlr;is,     lL'7. 

CIIAITKU    I. 

j'llSS.iil.iH.Y    OK   TMK    DK.ISTIVK    (il.ANliS    ((•ilNl'lu I.'.H 

N'liiiiial  ('llll(litioll^  of  ScMn'tiiiii.  iMi;  Nipiiii:il  Siiii'tinn  .it  H!ili\;i.  1-1:  Simii> 
timi  cil'  (Jii'itrii-  .luin'.    I.'.:.';   Tin'   liiti'stiii:i|  SiMirti'm-i.    III. 

CIIAI'TKK   1,1 

Till     Mkiiasisms    ok    I)i<;kstio\ Ill 

Miisticiiti Ill:    Deglutition.    II.':     TIk'    (';iiili:ii'    S|iliiiirti'i.    11*:    Vniiiitiii^'. 

WX 

ClIAl'TllK   Ml 

I'm;    Mm  iiAM.'^Ms   ok    l)n;t;.si-io\    (Cont'iii I.'>I 

Xlovciiiciits  III'  thr  Stiiniiiili.  l.'jl  :  I'li.'iuiiti'i  of  tin-  Movi'iin'iits.  I.'il  :  KtlVit 
of  till'  Stoiiinrli  Mnvi'iiii'iits  on  the  Knuil,  l.")l:  liiii|itvin;;  of  ilic  Stoiimrli, 
l.'jil:  I'miloil  Mf  thr  I'vlniii-  S|.liinrtri,  l.'iil:  Hiiti'  of  i:iii|it,viii'^  of  the  Stoiniicli. 
!.*>*:    liitliit'iH't'  nf    l';ii  jhit.i^if   ( 'ontiitioiiH  nti   till'    l\ih|it>  iM;x.    I'll':    'i:t>triM'jilt'r 

ostnlllV.     ml. 

(IIAI'TKK    I. Ill 

Till       .Mil  ll.\M.SMS     OK  |llOKST|o\      (  ( 'o\  1  'li  i .((J:; 

.MiiM'ini'iils    of    til.'  Iliti'stini'^.     Hiil :    MoM'Iiu'IiIs    hI     tin-    Sin;ill     Intotini'.     |li:i : 

MoM'iiii'iii-i  of  till'  l.tiiyi'  liitc.'itiiu'.  lilS;  Kll'i'il  iif  Cliiiiijil  ( •(iiolit inns  on  tin' 
Movt'tni'iits,    170. 

•  IIAI'TKK    I. IV 

lllMiKlt      .\M1       .SfrKTlTK |71 

lliinni'i-  Contiio-tioiis  ..f  stcnii.-oli,  171:  Koiiioti'  Klfi'i-tv  of  lluii;;i'i  Contrio'- 
tinns.  171:  lliiii;;rr  .liiiii.:;  Stnu.-iti.m.  I7.'i:  Cmtiol  .,f  tin'  lliiiiyoi  .\Ii','li;iiii>iii. 
4:ti. 

(•IIAI'TKK   I,V 

Iltoi  nr.MicA!,     I'Kon  s!<rs     u:-     Dioi  ^;;iii\' 1^5 

I)i;;i'stiiin  in  IIic  Stoimiili.  1*1:  I'limtious  of  i  In'  II  vilroilil.n  ir  Aiiil.  l•^-: 
Anioiiiil  anil  Niiin-c  of  tlio  .\iiil,  4SJ:  Ai-lion  of  l'i'|,>iii,  !*.->:  Cliitfin;;  ..f 
Milk  In  till-  Stiiinnrli,  4SS. 


will  niNTKNTS 

III  AITI.U   |.\  I  f,^,,^ 

HlMllllMIc    M      I'llMi  I  .SSI  ^^    lir     |)llil  .M  KIN      if'iiSl'lu IH'.I 

l>iU(-.!i..ii    ill    111!'    liili'Miiif^,    Isii;    riiiirinitii'    l>iui'sti..ii,    |>.!i;    Till'    llil.'.    In:;; 
ChiMiiUtiv  ..(■  Ilili',   liil. 

I'll  \i'T!:i{  i.\  II 
lUcriiirM,    iMtii  sriov    rv    riii     |\iisiim; \\i[, 

IllirtiTilil    l>i>!C.tlM|i   „f    I'mtrili.   .■,li|;    ll..tl|liMiii,  .Mi:l, 

PART  VI 

Till-:  i;.\(  KKTioN  ()|.'  I  |{im: 

I'll  AI'I'IK   I.VIII 

TllK    KX(  KITloN    III-     TulM      i   li\     H.    li.     I"l   Mil  I    , 5(17 

KtiiH'tiiiv  i.f  Kiilliin,  riiiT;  Mi'diiiiiisiii  i.f  tl„.  K\cn'ti..ii  ..l'  I'liiir,  ."In;  'I'l,, ,,,,,., 

Ill     Ki'iiiil    Kiiihtinn,  ."ill:    |)iiiictii'i4,   .'ils;    AMiiiiiiiiiiiriii.   :illi:    ImiIiu'I f    Ih,. 

\i-l-\iill«   Svsliiii    nil    llif    Siiirlimi    lit'    flilii'.    .'il'l. 

I'llAI'TKU   l,l\ 
TiiK  Amihm-.  riiMi'iisiT|.i\   AM,  Cii.'.uAi  rm  ot  riii:  l'i:isi    -  |i\    |{.  c.   I'l  u;,  i  ,     .  .-,j| 
Aiiiiiiint,  .-iL':;:   S|.iiili,-  {li:niiv,   .-.'J:    |»,.|,r,  K,.i,,ii  ,ii'    Kinviii-    IV, int.  r,-2:'.:    K,- 
.•I,  til, II,  ."L'l;    .'<.,li.l  r.,iistitiii'iits.  ."ij.'i. 


I'AKT  Vll 
.MKTAItol.lS.M 

rilAlTKK  l..\ 

\ln  AiKii.isM ; 

KiiiM-v  i:<il:iii,c,  .-,:;,-,:  .M.th.i.U  f,„-  M,.;„„riiij,  |-:ii,.it.'y  Out|,iit.  .-,:ii;.  .\,,iiii;,l 
Viiliics.  .-liis;  hitlii,.,,,.,.  ,,f  \^,.  ,,,,,1  Sr>,  .-11:  IiiMiiiii,,,  ,,r  1)i„.mm's  .",  |J  : 
Till'  M:,trii;il  lt:il.-inr,.  ,,f  till-  IJ-mIv.  ."I:::  M,.il„,U  f,,i  .\l,.;,Mirin:;  Oiil|„it.  .-,|;;: 
<':il,iihiti..ii  i.r  till'  Ki'siilis,  .•,11. 


CIIAI'TKK   1,\[ 


Tim    I'AKiins    llM.wri: 


I{i's|,iriii,,ry  (^ii.,ti.'iil.  .-)I7;  liillinii,-,.  „(  |)i,.t.  ,-,(7;  Iml,,,.,,,,.  ,,C  .Mrtiil„,[i,sMi. 
•'il!':  Miiuhitu.li'  oT  til,.  I{,'s|,inii,,n  i:\,-liii,i^,..  .-,.-,11:  iml, ,,.,,,.,.  ,,f  i;,„|,  ■p,,,,,. 
|„'i;itiiri'.  .-i.-il. 

rilAITKK   I..\ll 

.\  I'l.iMCAj.  Ml  111,, I,  hm;   I)i:tii:mimn(;  thi;  Hi  si'n:.\T,,i:v    l-.\,  ii  kM:i    is    Man   (J',y 

H.    li.     IT.Ai;ci:) 

Till'  Viilvcs.  .-i.-i.-i;  'ri,<s,,t  S|,iri,ni,-t.'m,  .'i.'ir, ;  Dmiyl;,.,  |!;,i;.  r,r,S:  Iliii.liiiii'  Gas- 
■ill.-llysi-i    \|,[,;il;itili.  .-..-i!!;  I'lili'lllitt  i.ills.  ."(iL'. 


\w 


ruVIINIs  M\ 

('II  vniK    I.MII  fAiiR 

Si  MIVMIIIV ."ititi 

UMirlimi  'I  NiliiiuiMi.  ."illil;  l-.Mii;i.\  Uiil  |  ul,  .'itl^  ;  \iliii;.rh..ii«  Ml  iiil..i|iii«,  ."ills  ; 
IlkiTi'tiuii  iif  l'iiiihr>,  ,'il!i ;  I.M'Mliiiii  III  Siil|iliiii.  ."iii'.i;  Nmiiiiil  Mi'tiiliiili>iii, 
.'7'*;    \  it  lii;;rliittl"«    lii|'iililil  iiiiii,  .')7I;    l'l"iriii   S|ijhi-i-,   ,'i7I. 

•  11  MTi  i;  i.\l\ 

\i  i mi io\    \Mi  (ii;it\\  III     ....         ri7( 

Till'  I'liii'l  riii'tiii  Hi  (ii.iutli.  :.;i.  It.  Iiitinii.liip  III'  Pint'  ill.  Id  <iiiiulh  ;>li.| 
MiuiilriiiiniT  iif  l.il'.i,  .■i7l. 

I'll  M'I'I.U    I.W 

Si  iiililiiN     wii    IIkiimiii     iChsi'ii- ,■,>:; 

|{i'liitiiiii>lii|i    I.I'    Cm  i...li\.liiiti'<    .■in. I     l':i'«    In    ili.iulh,    ."is:; ;     Ai'ic'««i.m      Y' ' 

I'llrt.iK.   ..I     \  illllliipil-.   .'lO  I  ;     liilllii.|lslii|.   ..I     I11..1  :;,-illii-   SlllN.   .'iMl. 

•11  M'l'l.i;    I.W  I 

I'll  It  III  s -,^S 

l':iliilii.     I{..||llilr Ills,    ."iss  ;     Till.     I'n.tilli      I{|  i|il  i  I  I'tinlil  ,     ."I'.iM:       \iii«,.i|i       I' I 

l'";iiinM.  :,■.<:];    |)i^i  .tii.iiitv    nii.l    I'.'ihitMl  ilil>.  .-'.1:;. 

(  II  AI'TLK   I.W  II 

Till.   .\Ii.im:iii.i-.m    m     l'i:iiiii\- .Ill,-, 

Iiitr.i.|i:.|iii\,   .•,|i.-,:    rlii-iui^li  i    III'    I'l.iti.iii    iin.l    III'    111,.    Aniiiiii     \.il>,   ."i'.i7. 

('iiAi-ri:i!  I.W  I' 

I'm       .MkIM:iiLI.-.M     lit      I'l.i.lM^      (I'.iST'ii,        .  IjllC 

.\iiilii.i   Aii.i-i  in   ill.     Ill 1   :im|   Ti».,iii.».  Cm;;    |  ,        ,,|    i),,.    Atiiiii.i   Aii.|-i.   lilo. 

I'll  MTlli    l.\l.\ 

Till     Ml.T.MIiil.lNM    OK    I'UIITMN     .('OM   'h, ,;|.; 

illiil  I'li.iliirN  ..|'  I'luti'iii  Mit:il.iili,Mi.  iW.-,;  ri,;i  ,.,1111  Aiiiiiiiiiiiii.  >■''•:  In 
thii.||i<.  Ill'  AriiiiKi.i  nil  Aiiiiiiiiiii;i  nil':!  Hntin,  illi'.;  I  iiiliiriiii.  ni'  l.ivi  i  .11  Am 
iiiiiiii;i  iin.^i    Kiitiii.  ilir.    I'i.||ii.i r   ()i-..|ii..,   <ll>;   <'liiiii-;il   OI.scimiI  iniii.   liUH. 

I'llAI'TI.IC    l.\.\ 

Tin.    .\Il  T.Minl  1>V1    111     I'llOTHV    .  < 'DM' 'li ,;.... 

I'li'iitiiii'  an. I  rM.,,tiiiiiii'.  I'll'l';  lis.inl  iai  r|„.„ii,;,|  |.';i,.|..  iijn.  M,.|„|„,||^|„^ 
I'lL'l;    liilliii-ii-i.  ..1    K..1..I,    \t:i.,  ami  .«,.x,  Ci'l:  Oii-in  ,.r  CiLaiin..  aii.l  Ciraliiiii,!.. 

ii:;i:. 

i   IIAl'Ti;!!    I..\\l 

Till.    .\ll.T.\lai|.l>\I    ()!■     PUOII.IS     iI'llNl'lii ,;..,, 

riiili'ti'iiiiiiH-il  .Nitiiiticii  aii.l  |)it..\i,:ili..ii  ('.ini|...iiii.N.  r.J:i ;  Ijlirn.:,!  Sii!|.lialr-i 
ami   CliiiiTiiii.-itcs.  C,?,-2. 

"llAI'TKIf    I.. WII 

I'kic  Ann  AMI  nu;   I'l  i:im.    n.ii.us ,;■;! 

Clicmlral    Nature    i'!'    tin-    IViriia's.    C::!:    Ch.  lai.-i!    V-,t.ii.-    ••!'    i!:.-    S;:! -lam- 
(.•iintaiiiiiiy;  I'liritK-  ami   IVrimiilii:.'  pji^rs.  i;:;7 :   llistm*   of  Nm-liii-  A.i.l   in  tin- 
Animal    limly,  Ulis;    ]!alam-r  l.rlwrni    Intaki'  iiii.l   ()iit|.iit   iif   riuiin.  .Snlistami-- 
iimliT    Vaiiniis   riiysi.il.i_^i..   an.l    l'atliii|iii.'ii.   ('(imliliiins,   (111. 


^^  1  us  1 1  M.» 

'II  MTl.lt   I, Will  ,.„., 

I'lill       Aim     \\l>     ll'l      I'llIM       lllllHls     iCiNl'ln  ,[|; 

**•»"'■'■     "•'     KM.I..1J I„     I'llMll.,,    lil.i;      liillll.n.r     ..I      \,i,i,„l<     ni».i"l"ul''    < '"» 

.lilji.n..    „f    Dnm-,    iih.l    „i     hi,,.,,*.    ,,„    t|„.    l^:,,,}.,;.,. I,    rii,ii,i.|    K\.-r..|in.i. 

•ilT:   I'lii    A, 1,1  ,,r   III I    ,;(, 

<  II  VI'Tl.li   l.\\l\ 

Mll>ll.M*,M     ,„       nil       ri|;M..HM.Iail> ,1-,^. 

•  •|i|.ji.ilv    ,.r  11...    I1...U    t,,    \,.i.„lh,i..  r;,,l,„|,,, I, ,„,.,,  ,;.-,j;    A«Mii.ilMti..H    l.iniiu. 
"•'■-:  '^"'"liii l.iiiMi..  i;.-.i;   |>,;...,ii..h  iii„i  AI«i.i|.iiMh,  iMii;  siimii   I^.m'I  in 

''"'     '" '•    '''■''';     ^'ll'"'    "I      lll""'l     KxilMliMlltilMlH    ill     l>illUMII..i«    ,.l      IliaUlr,,    im!>: 

"•■'"''"""'"I'    l'"t»..'i,    I'.l I    Si^ar    mi. I    tli,.   O.-nii  i.ii.c   nl'    (Ihr.i.iiriii.   liiWi. 

<  II  M'TKK  I.V.W 

\liTMi..i  i.-\i     .It     r >i;iiiimm.i:mks     (r..\i-|,| ,j,j_i 

l-'ul..     Ill       \l,,i.i|.,.,l     lllii,. (illli-,.li|.My,.MCNi-.     IlliU;      M.,l;i^..     iiT     .Sn;;,,!,     (ii;;;; 

SiHiiiis   ,.|-  lllv'i^i'i.   'iilj;    (ilii.niii„.^,.|ir.i.    in    \,,,iii;,|    AiMiiihN,   iln:. 
<  IIAITKK    I.WVI 

Ml   I  Ml'.l,l-.M      III       I  Ml       I'M: Mil,' Ml.-.       iriiM'lii        . ,;,;,, 

r.ilf    I.I    lilM-..L;rli,    lir,:i:     li,.u,i|:;ti I     ;!„.    |;l 1    Si,;,,,,.    i,,.^,.\     ,;:]■.     Xrn,. 

'■mill. Pi    (III. I    l;x|.riliiM  ntiil    l)iiil.|.|.N,    iirj;     NiiMMis    liiiilirt,.,    in    X|„„,    liTI: 

""" •••■Iltr.il     Mll.l      P,.,,„;„ I       IlilllTl,.,,     Iir.l;      l|ili/„|i„,|      ,   ,■     (i|,„..,„.     i,', 

Tis,.!!,.,.  i;;7:    Hrl.ili r   tl„.    l'!,,,,!,.,,,  t..   Sh-mi    .\l..tiil.,,li,i,i.  117s;    |)i:,l.,.|,.s 

.'III. I  III.-  |lii.il,.«  (II I-    .;;s;    |ii,,|„.|i..   A.i.l.'Ms  i.r   K.i.„iH.  lis:;;   ,Sii,r»i.ii..M 

Tl.'.lthi.llt.  IIHI. 

<iiai'Ti:k  f.x.wfi 

I'm    Miimioiism ,.m. 

rh..|||i,tll     ..r     l':ilH      .S„|„|;,ll.-,.,.    IISC;      |l|^..»|i,,„     ,,|      r,,u.     iliMI;     A  l.«.l  |.t  iull     ..C 

Tills.  li'.H. 

'•|lAfTi:i!   I.WMII 

I'vl      MllMl.ilISM      |I'ii\t'!ii ,;,„; 

'■'■'•     "'      "I I.    '''■"•■     M. -III. "Is    i.r     Dl'IinniMillliili,    IiIM;  ;     \  ;ui;iti,,||s    ill     III 1     |'.,I, 

ii!'7;    |li.|.ii|    Kilt,   71111;    Pill    ill   tlii>  I.iv.T.  711I, 

•11  \l>TI;l(    lAXIX 

I'm    .MllMl.ilISM    ii'iisr'iii .   . 

''""'"'■'' '     '■■••'".^     \'''l    "'I'    "I    <'llll.|||l^.llllt.■.    7h7;    .\l..t I    In    W|,i,.|i   tl„. 

I'mtv   .\.iil   i..  I'.nikiii   ll.iHi!.   7111.. 

lllArTIK  I.WX 

i'.i\Ti:iii.  Ill    r.iiiiv    'I'l  Mi'i  ii.vii  i:k    km,    |.'i  vi  i; -,  , 

V!irlnli..iis  ill   l'.,„lv  ■|'..|Mi,..|,,tiii.-,  711:    r.i.t.iis  ill   .Miiiiiiiiiiiiii^  tin-  li.i.K    T 

(•.■nitiii...  71.-.:  i'.,Mti..l  „f  T.'ni|...,:,(„,..,  7l!i:    |.-,.v.-i-.  7l'l  :  Caiisi.s.  7l'l  ;  Chanyi.s 
ill   III.'   r..i.lv    li.niM-    K,.Mi-.   7-;;:    llrat  i.'Huliitin;;   ('..|itiM.  7-.-):    Simiili.iin,,.  ,.|' 

r.'MT,  7L'ii. 


i;i;<) 


I'AUT  Mil 
TIIK  KMM»t  KINK  n|{(i.\NS.  t»|{   |»l  iT|,|;ss  OLWDS 

•  11  \ITI.K   I.X.WI  ,.»,,, 

I'm      i:\INMI.I\t    (diCWS.    IMi    III  1    II  IKS    111. AMIS •■», 

Mi'IIiimU   nf    liiM'^tiuiitinii;    T.'.n ;    Ailfiiiil   tihin.l,    Til;    C.itis,    Ml;    Mfiliillii, 

".'.J;    A.liiiiiiU'.-idiiii.   ".'.:'.:   Hui'iiii'hil    Kxtiml-..    7  11:    rin»Milnyji'    A.tlon,   7:i|. 

(  IIAITI.K   I.WXII 

AUKINAI,      (ll.WII      iCllNl'l ...^ 

\iiiiiiti"ii«    ill    I'hv.iiili.uii'    A.IInIIv,    T.!":     A-.»iiunj.r    tlir    Kpiiii'ipln  ini'    «inili'iit 

111'    till-    llliiihl,    r;*;     K|'liii'|<liriiii'    rmiti'iit    ..I'    Ihr     III I.    7;'.!t ;     Aiil"iii  j.'ctioii 

Mi'lliiiil,   7t:i;    \.lrin!,lriniii,   7l."i;    A^-."iiiili i    iIm'    A.lniiiil   witli    il'li.i    |.r 

ilmlillc   (»||.'1IIM,    7  |l'. 

<  IIAITI.lt   I.XXXIII 

TuMii'iii    \MP    I'\i;aiiim;iiiii  (ii  \mis -^^t 

Hlnicliirii!    Iti'l:iii..i,,|ii|..   7Hi:    -riniLi,!   lilnihl,    7:iii:    i 'iin.litinii   oi    (iliin.l,    7Aip: 

KviMiiiH.iiiiil    Tli.n..i.|.'.i K.    7.">i':    Ki-mni^"    ..i'    iIk'    Tlnn.i.l.    7."i:! :    Ki'liitimi 

Hith  Othii    liihlii.iiiir  (li^;,ii«.   7.*i7:    I'liiiitliM.ii.N,    7.'><» :    |:\|h'i  inn  nliil    I'lniilln 
■■'•■itniin  .  7,-,s;    |{.  Iiitii.ii«lii|.  with  Othn    Ijili.,  i  in,'  OiyiUK,   7(1|. 

<  IIAITI.I1    l,XXXI\ 

I'm  iiAia     lli>iiv -,.., 

.'<liii(Hiriil   K<'lati^Mi-lil|.«.   7il:.';    I'uim  ii,.ii-.    7HI:   (liniinl   r|,;,,ii.fi  li^ti.  s.    771: 

Krlutiuli>lil|i    Hill,    UlliiT    Kll'liulilif   (It;;!!!!",    77.:. 

•  IIAITKK   I.XXXV 

Till.    I'l.MAI.    lil.A.Mi    A.Mi     nil     UiiSAhs 77(i 

riiifiil  Cliiiiil,  77ti;   (ioiKhli  ,11    till'  (Miiiiiili\.-  ()i-iiii»,   771'.;    (..ni'iativf  filiiii.l- 
"f  till'  Mall',  77<1;  ("■niiali\r  ()i;,aii>.  i.f  llir  Kniiali'.  77s. 


I'Airr  i.\ 

Till-:  <i:.\TI{.\l,  NKIiVdls  SVSTKM 


n  I 


CIIAI'TKK  I.X.XXVX 

TllK     KVUI.fTKlx    UK    TIIK     XKIiVnl  s     SV.STKM 


.  :si 


<llAl'Ti:i{   I.XXXV  II 

I'Knri.inii.s   iiy    Mai  ii    I'aim    oi     imi.   IJi  i  i,i a    Am ;^^ 

K.'.v|,i,„.  7ss;    Kpinitir  aii.l    l'i„t„|,ail,i.-  K..,-,,,,,,,.    7'.mi;    |v,„liaiiti..^     r  tl,.'    ' 
.Si|.aiato  KciisaticiiiN.  7!'l  ;  T.iii|..Taiiin'.  7!i|;  T.mi.  Ii.  7!i:;;   Pain,  711.-,. 

•  IIAI'TKR  I.XXWIII 

Tm:    l'liO,.KKT.K.S  OK    Ka<I1    I'Ala-  .if   THK   KkKI.K.V    Ak.'    (CintV,  7;r, 

■I'll..   N..n..  NVlunrk,  7:m1;   NVt«,„k  ..n  ,>.ikin   X.tv.v,   7!m;:    TI„.  SMi.,|.si<'  T'.7' 
Til..  N.TM.  ('.•11.  7;'|.;  Tl...  Ii,;..rii,...liat,.  ..i    liit..i  nuii.ial   .\,.,i,,.„,  vnj. 


NMI  ("ONI  i:.\is 

<MI.\l'Ti;ii  I.WXIX 

liMiJMs  HI-    Jill:  M'ISM.    Ammai     -Ml  sri\M    Shock so;; 

S|iini(l   Sh.M-k    In    l.;il.(.i:ili.i  \    .\iiiMi:il>,   m'.;  ;    .S|,ii,;,l   ,sli,.,k   In    M;in,  Mill;   (•.•iiisi- 
n|'  S|.|n;il    Sji.Hli.   M17. 

(■|i\i'Ti:i{  .\c 

I'iU^iHi  .1,11     I'K.ii-i  Mil -,   ..I      Mil     .siMi'i  I     1(1  I  I  1  \     \i:<- sii;) 

l.llti  ill      i'lli...!,     Sli;i;     (;|;|,|i||;;     ,,i      |  ||  1  I'll  nil  \  .     Vll'.l;      .\  It  IM    r  |1 1'.t .     ^lO;     Sll  llllllilt  Ic  III . 

^|i';    h  ii'M  imMIIIv    ,,r   III,,    hiii.ti r   I'mi.lii.tioii,    "-In:    l{iri;i,iun     I'l'iiii.l, 

»l  I  ;    Sill,  rsvix..    Il.^iiii  iMlloil.    '<|.;. 

<  IIAI'TI.H    M'l 

K'  I  ll'Kdi    M,    l\MI:V\llll\ \\\ 

Ur.  i|i.i.:il    III!, il. ill. .11.   .-II;    A.li I    Simliiiiii..  :ni,|   Trimiiiv   'rn.\in,  s|!'. 

I'llAI'Tl.lt    XCll 

|\iii:MNo.\      Am.im,      IJiiiim- SJl 

lii!'^i:.ti..ii     ..!     Alii, ..I     K,-ll,.\,.,.    sJJ:     Inl<'i^i:iti..n    >•(    Aiit:i-..nl>th'    |{cM,'x,'s, 

^L'l;  Oili.T    I'.-i.i...,  Wliirl;    K,!.  uniii,.  (i...||,:in.v    ..t    I'lnnl   C.i n    I'atli,  >*L' I ; 

Ii  i;i.lliti..ii.   ^L'l'.. 

(HAITI. 1;  .\ciii 

Till    Ti  \i"i\  .1 1  i,ii>  :  Si  \.-,.ii,\    l'\  1  .i\'.  \\  >  i\   Sn  N  \j.  ( '.ii;ii vjv 

Tl'i     'I'l-li.! l,'lK>.    »L'^;      \IV.  l;-lil    S|.il'iil     I'.itliuiiv.:.    ^.Iii. 

<  IIAI'Ti:i{    XC|\ 

I  J  i  I.,    i.-    .•!■     I.Mi  i.lMI    \  :   M      I.I   --:..\s    i.|      \'  \  K  h  .1    ,    I '  M,  1  -    .  .|      1  1 1 1.    .\  i  la  1  il    >~    S\  s  11   M        .^.'l.') 

.\iit,Tlc.i    1{....1~.  ^.;.".:    l'..-l,.|i.ii    l;....i,.  >.:;,;;   Spin.il  i'mi.I.  ;,i,,|    I;,:,;:,  St, .111,  ■*.■;!,; 
.Mi.hillii,    v:;;,;    c.ii.Mi,,    (^)ii;i.|ii^.,  mil,:,.    v|(i;    |{,.,„.,i;i|    ,,|-    t!„.    C.n.l,,:,!    H,-inl- 

-|.|!,.H-,    s(,,. 

<iiAi'Ti;i;  xc\ 

ClMliKM      l.ni    MIZMIUN y  ( ;. 

Al.liitiiiii   ,.t'   tl:,.   Mut..r   C.I, !,■'-,    v|:;:    Sii.mil;,!  i ,h,.    M,,t..i'   Ciii,.,^,   ^|l: 

i'liiii.;il   ()t.-,.M;iti..ii^.    -t'.i. 

(■|iAi'Ti:i;  xc\i 

("l.ltrUKJI         l.n,    \l,l/\||l)\         iClNl'l.) S,",0 

S,.i!v,,ix    C.iii,.,^.  s,-,n;    S.'n-..  C.iil.-iv.   s.-,l:    A.-...-i!iti..ii    Ar,.;is,  v.'L'. 
(  IIAI'TKK    Xc\n 

ClNlll  I  Ins  M      ..,M,     I'm  .i\|.p  li.is  M       lIllll.M.-- S."itl 

(  ilAI'TI  1!    X('\lll 

III'. nil.     I'l    N,    l|,,NS    ,.1      I'll'      I'lili.lllM     1\     .\|\\;     Al'H\--l\ Mill 

I'^vi  !i'i|i.itli.'l"i;li  ;il    .\|.|.li.  nti'Mis.    m'.l'. 

<  HAI'I'KU    XI 'IX 

T'si  iKiNs    III      Tin:    ("I  Ki  nil. MM  S,r,." 

t,...ii;,/;,ti,iii    ..r    I'll,,   li.. II.    »,i7.    I  iHiii.isri  iI...,|    l.\iir|.:iti..n.   Hil'i;    ('linniil    ()l, 
Ml  \nti..iis.  K7ii. 


■'a 

is 


t 


a 


CONTKNTS  XXm 

illAl'TKK  C 

Till    '  I  III  111  1,1.1  M   .\Mi  nil:  SK.MiiiKci  i.Mi  Canals;  Fi'.nciihnai.  Tk.sts     ....  S7," 

Mci\(Miiriits    iiiiil  tlu"    Scniicirciiliir   Cjiiijils,    ^'o. 


ANS'M-iiitiuii    l.i'twivM    til"    Kvi 


(  M.Vl'TKu  CI 

Till:    .\Mii.\oMic    Ni:i;v<iis    S\.stk\i S7i 

(ii-tii'iiil   ri;iii  (if  Cdnstniiti H77;   Tlii.iiii-icnluinliiir  Outtliiw,  in    Sjnipatlictic 

Sv.stiMii  I'ldpiT,  SSO;  liiilliii«ic'i!il  Outflow,  r,r  till'  I'iiias.vnipiitlictic  System,  SSLI; 
Axiiti  Hcllfxcs,  ^s.",;  KiMicliiiiis  of  Aiitiirii.ini.-  Ncrvi's,  VSI;  Atrcrmt  I'IImts  (if 
the  .\iitiiii(iiiiii'  System,  KS5. 


: 


1  3 


1) 


t  mi;^  I  til 

9  ^B     Ml 

^  Pi    In 

in:  mm--        t^MM 


ill 


ILLUSTRATIONS 

'"••  I'AM 

1.  !)i;it,'r;i!ii    i.l'    c.^iiinnu'ti'r - 

-.  llciMiitipiritr 

.'..  I'lasiiiiilvsis  in  rills   iKiiii  TrailcM'aiiti;i   ilix-nlor i| 

I.  .\|.|.:n;iliis   f,,r  iHi'Msiirrnirnt  of   the  .lf|.ics~i f    rnr/iiiL;    |inim    ,,r   >..|iiti..ii      .  II 

•".  I>i:i;,'r; f    idinliji-l  ivil.v    cclU Is 

i>.  Wliciilslciii'    liiiil^c   f(ir   till'  iiicii-diri'iiiciil    (if   ..Ici-lrii-    riT<isliiiii'.- Is 

7.  |)i:i^i:iiii   t.i  slmu    tv|i,.  i,f  i-lcctniilrs  iis.'.l    in   stii.lviiii;  rl,'.-ti.ii;i,tiM     fiivi,.     .      .  :;ii 

:■.  <'li:irl    III    tints    ;is    iisnl    in    riilm  iiiii<i  rir    iiii'n>nH'nii>nt    nl     Minn    .nnrrntrnt  imi. 

ifi.lnl     l'l;,ti-. ■;!   ■ 

•<.    Ili.-ij;i,-iiii  iif  iil'i'.-natiis   fm-  ihr   niiaNiininrnt   i,f  ilir    ||  inn   iiini-t-nl  rat  inn      .      .     :\\ 

III.    Oia^'iiini   iif  appaiatiis    fnr-  satuialinu    M I   ami    plasma    «itli   i'\|iiii'il   air      .      t.'. 

II.    \an    SIvlir's    aii|'iiialus    fm-    ninasiiiiiiu    ilm    ( •<).  inniliinin-    jiium-i    nf    M I     in 

liliiii.l    plasma .|.( 

IL'.    rilraiiii.iMsiiipn     ,,|it    tvpri     tm     ilic    i.\aminalinn    nf    mllniilnl    snlininns     .      .     ."ii' 
l:;     Tn    slii.u    ijiiru-inii    inin    M,.|;itin    I.I     a    i-rvslallniil    stain,    ami    llu-    min.liirnsii.n 

nl'    a     i-nllni.l  stain ,",:; 

II.    Diagram    finm    W.   Oslwald   slinuin-    tin-   n'lati\i-   sizr   nf    varinns    paitiilrs   ami 

i-nllniilai     ilisprrsniils    i-nnipafi'il     «illi     :i     ml     I,; |     ,       niis,li.    atnl     ati 

antlcrax      liacillns ......     ."1 

l-'i.  ''a|'il|aiy    .-inalvsis    nf    ii.llniils .-;,; 

Ii'i.    IHayiani   tn  slinw  stiui'tnm  nf   m.|s i;| 

ir.    |)ia;;iatn    tn    illnstiat.'    sn.rfaii'    ti-nsinn (■,,( 

!•>.   'I'laiilin's  stalaL:mnni('tiT li,-, 

l!i.    Hianinin  nf  till'   ^-rapliii-  niaunlnni.'ti'f |(iii 

-II.  '■nai^nliimrtcr ^  ID 

-I.    .XtiiTiiiy  inaiiiiim'tni   ami  si;.'nal  tn:i;;iirt,  arr.-inLTi'l   fur  rninnliny   llm   nn.'in  ar 

tiiial  Monil  picssiirn  In  a  lalmr.atdiv  nxjicTinii'iit Il'-l 

-L'.   'riic   arti'iial    lilnml    |,irssini'   ri'i-nvilnl   wltli    a    int'icnry    ma ti'i     ;  Inwi-i    lia'- 

iii;;i  .-linn;;  uiili  .a  traiinu  nf  tin-  iv-pivatniv   mnviiiiiMi    nf  tin-  tlinia\      .    I.'.' 

-'■',.    Iliiiililii's    s|iiii,^    Tnaniinu'trr Ijii 

-'■   -\iliTi.-il    pii'ssnH"   ti iili'il    liy   a    spiinu    ma tnf IJi; 

-."i.    [>ia;;iani    I.;, -ml    on    ."xpci'linnits    nn    ili.ys    tn    slinw    tlm    svslnli,-.     ".aslnlii-    ami 

ini'an  I'lnoil  iircssnrcs  at   ilKTcr.oil    parts  nf  tli,.  rirciilatciy   system     .     .    l-'7 

->''.  .Vpparatiis  fm-  measuring-  tlio  arti'ii.al  Mnml   prnssiirn  in  niati 120 

-7.    l^lTi'i't  nf  iMittiiij;  tlip  vatriis  iiorvc  im  till'  artcri;il  Iilnn.l  prcssnri' i:i.". 

-•^.    KfVi'.t    nf   stiiiiiilaiin^    tlm    pni  lplmi;il   imi.I    nf    tlm    i-i-lit    v:i:;iis    n-i    tlm    ait.-iial 

Mnnil    |ii('ssiivi'        |;;i; 

J'.i.    i:irivl    ,,|-    stinnilatimi    .,(   tlin    l.-ft    -plan.lmi.-    mivr   nn    tin'    aiti'lial    lil I    pus 

^iiii' i:;: 

'■'■I'.   'I'hi'  .iTiit  nf  r.'ipiil  ami  sinu    licnnirrliaui-  nn   tin'  arti'iial   lilnml   pn's.siiii'      .     .    |:;.S 

■!1.    Di.'iyr.'llll    nt'    nxin'linnnl     tn    slli.»     tliat     III,,    ijiastnli..    i.r.'ssnr.'    .lopi'ii'ls    "!!    tin' 

ilastii-ily    nf    111,.    \,.ss,.|    Mall 1  C! 

■'.-'.    Hiauiani    nf     Wi::^i.r..  •    .,|.iii..'i|     m.-innnn'ti  i 1  in 

x.w 


XXVl  ir.l.rsTKM  IONS 

^1.!.  Ujitiral    ri'fiinls    «(    liiti.-mni  i  icuhir    jiii~>iiii' 117 

;:i.   Sii|.iMirii|..isc',|  |.iis.iiir  .•iir\i>  :illii    l.riii^  i;i:i.hi:it.- I I  I'.i 

::.',.   \'..n    Flunk's   m.-ixiiiiiil    ;iii.|    iiiiiiiiuiil    v.ilvc,    wliii-li    is    |ihi,T,l    in    tlu-    i-.mhso    of 

111''    liil"'    liiIiMTii    Infill    Mii'l    iii.ii'iiiy    iiiiinMinc'l.r I'l-J 

■  '•I'.    I'i:i;;r;iiii    In    sli,,u    tli.'    |".sili.ihs    ,.t'    th,.    .iirdiac    \;il\,>s  ].-,;, 

.;:.    Iii:i-i;,in     -h.miny     ,1,,.     |.,,viti,,n     <■!     Il :ii.li:i<'    ,Ikiiii1„  r ;    ;,n,l     x.-ihrs    .liirill? 

I .v~t..||.    ;ni.|     .hiiirii;    til,.    s|,|,iiiii.-    p,.r-i...l ].-)i; 

-'<.    I'ili .  tn.|.|ii.ri,..^i;iifN    .■iIoiil;    witii    iiil  i-.i\ .  ii!  i  i.iil:i  r    |.r.-.Mn.-    i-iii\.s    |'i<,iri    ilir<'i' 

ilillrrciii      rvi..!  iiiiiiiis         1i-|i| 

■"'.'.  On.-  iMiin  .,t'  ;q.|.:ir.()iis  t,.i  i, .,,,,.!,,„  ir.i.inu-  l'i"Mi  :in  ,N,is<-.l  h.'art  .  .  \i\:. 
|i'.    \'..l;iiiir    .IHA..    .,r    v.iitri,  !,..    ,.(   ,:,t    >l,,u,i-    ,iha  ,        in    :<    liruii   Inili.'    |mt  fusion 

l"''l''i'';i'i"'i        lC,;i 

n.    I|i;irl    :iii<i   c-ir.ii.'ir   lun.s  i,i'    I.imiiiIiis    |.i.1v].1li-iiiiis i;;; 

I-'.    llrMrt  l.l.ick    [.r.iiliii-.-.|    l,y    Mp|.|\iiiu'   .■iiiiiip i;:, 

l:,    Tl.-li'ill;:    of    i-f.Illlili-tii.M    ,,|     \,  nlri.l,  .    ^lii.uilii:    tin-    .  IV.rt     i.l'    tlir     lo,;il     .-ipiili- 

caticin   iif   li.'al    t,.   t[i,.   Mini.  I. ■ j;.-; 

II.  I'ro-  li,>:irt  stn.wiii;;  Ih,  pnsiii,,,;  ,,r  i|.,-  lii^l  mm. I  ~,  .-..ii.!  liu'Mtiins  ,,f  StMimiiis  ITf, 
I.'..    lifTi.ts   ,.(   sliiiiiili    i.r    iiM-icMsinu    -iifii:;!!,    ,11,    sk,l..t;,[    Mil. I    ,:ir.|iM.'    mils,'!,'    to 

lllustrMt,    till'  "mII  ,.r  M,.lliiiiLr"  prill,  ipl,.  in   th..  ImHit      ...  .      .177 

||'..   Til,'  .  IT.i'ts  ,,!'  sii,-c,ssi\,.  stiniuli  ,11,   vU,.),.tMl  Mini  ,;ir.liM,'   inns,!,.   :,,  ,,ii,,w   tlio 

IT,.niiii.n,-,-   ,if   111,-  stMir,Ms,     pli,  ihhihtici ■   tr,.)ip,.,   in    tin-   hir.-r     .     .   17S 

17.   Till'  .ir.ils  ..r   sii.-.i'ssiv,.    .tinniii   ,in,l   ,ii'  tctMiii/in-    stimnli    ,.ii    ^krli'tnl    nms,-l,> 

.■mil     r.'ir,liMi'     mils,  1,' ^-f) 

1-.    M^.-u'iiniis  i.f  frill's   x.iilii.l.',   ,i,,,vvii|.^   ,  |V,.,-t    ,.f  ,\,iiMi  i..ii    l.y    liri'Mk    iniliii-- 

li.'ll    ^ll..,-ks    .Ml     \, 111,1. IS    I  I,, Ml, '111-    ..I    ill.'    .Mi-.li: mU' ]S(i 

1'.'.     lI'Mrl      ,.f     t,.rt.ii>,'     M~     -llsp.li.l.  ,| 1^;; 

.10.    I)iss,.,.ti,,n  .r  Innrt    I.,  slmu    m in  i.nlin  .'lit  ri,iilM '    Inin.ll,' 1,8| 

•"'■    I'li.'li.;.'rMpli  i.t'  1 1,1  ,,f  111,.  Miiririilnvfiifri,iil;,r  l.iiinll,.  mm. I   il>   rMinili.'Mtinns. 

,..ii«liii. •!.■.!    fr,.ni    ,ii-.s,.,-ri,,iis   i,f   111,.   Iirinl IS4 

''-.    niii'.;rMiii    ,,r   MM    Miiri.  li'   slinwin^'    tlin   iirran^'.'iii,  >.l    nf    Hi,.   iiiii-,li>    l.tiinN:    tlip 

i"i iitrMli,.n    p..iTit:    Mii.l    til tlini'   nf    ili,.    ii,i,I,. ISfi 

.'.".   l>iM:;iMiii    f..    sIm.vv    fill'    ;,'i'ni'i.-il    rnnnfi.':iti,Mis    ,,f    tin'    fnn.Iin.tiii^    tissn,'    in    fin- 

lii'Mrf     .if     till'     niMninnil -[.sn 

.'.1.    I'lM^'iMii!   til  illiistrMt,-  III,    ,|,'M.|,ipiin.nt  an.)  spri'M.I  of  fin'  \\:iv,'  of  n.'^'ativity 

in  n  strip  of  inii-.l,.  .  ,iir.-iri/,Ml  sMrloriiis'l  ulnii  si  iinulat.'.l  at  tli,.  on, I  .  ISs 
.'."■..  Siiniill.iii,  ..lis  |.lr,tr,.,'Mr.li.,;;r;iins  to  sliin\  tin'  iMiisi'  for  ('\trinsii'  . 1, lie. t  ions  IPn 
."il.    ItiiiLTani  of  ,  \p,.riiii,iit  I,y   I.,  wis  sliowinu  li,,.   linios  at   uliiili   ttir  <\,'itMli..n 

\MiM'  Mpp.'.-ii.'.l  on  till'  front   of  tli..  lii'int  194 

.".7.    |iia;;rMni   of   1 'li.-iin  .  mh 's   .Iroiiio.rrapli 2011 

."iS.  DiM^iMii!  t.i  -how  I  lin.ipli'  .,r  I'it.'t's  tiil'i's  f,,i  iin'MsiiiiiiL.'  (.Ini'lfy  pulse  .  .  2ni 
I'lii.    Dinl-o.ni 's   spl.i -ii|,.^iMpli 201 

t'.l.     I'llK..     fl:|,i|l;;      ■-pll\  unio^'IMIn  .     InK.ll     l.y    spliyo  nio;;!  ;,  pli 202 

I'.J.    Form-   ..f   MppMrritn-    for   nn'Msiir,  noiit    of   Moo, I    \-,-l,,i-ii  io>: 207 

•''■'■.    I'l.tliysr     ^TMpli    for    r lin^    \,.Iiimio   .  lianc.-s    in    tin'  IimioI    mii'I    forrarin      .  210 

•'■i.   .''iiniullanpoii-    !rM,'iiit.'s    fn.ni    ;iiui.l,'    mimI    vontricio    of   tiirtl.'s    liorirt      .      .      .  21'' 

>'•'.    Ffr.-t    ■■f    Ml-lls    .!in:!llMti-ii    ,-.,:    hi-A-.t    ,.f    ti;i-!!,>  21'' 

Ifl.   Trillins:    'o    slmw    tliMt     \m_'iis    -'ininlMtioii     mm     .liiiiinisli     t  rMiisniission     fi..ni 

anriili's   to    v.'iit  ri.'li^s 21P 


II.I.ISTKATKINS  XXvii 

!'<'■  r.\(ii; 

|17.  'I'i;iiirn;    l.i    shiuv    lli;it     \;i;;ii>    >l  irnuhiMnii     n  .ix     l.-nililiili'    1 1  iiri-iin^iun     I'ldiii 

iiiiiii'h's   tu    \riil  I  iilcs ^:;ii 

li-.    I)ii4;rtuii   tip   ^ll.p\^    llir   iiihiuiiti jT   tl,r   ImmiI    in   llir    I'm.;;   ui    turllf.      iCc.l.ir 

I'lati'.  , '^'ji 

i;'.'.    I'rc.;;   Ii.-:irl    linriiij.'   vli,,«ir|.4    ilu-   iii'tiuii   of   uici.li'ir ■     .      .     .  2Jti 

:<>.    Silli'MiMlii-     r.|.rr^rlil;itiu|i     c,|-     till'     il l\:iliiiM     III      till'     l»;ill      '■!'     Illr     lii:l  nilll.'l  I. 

p  <■"!,, I-    I'hilr,  1 Jj,; 

;i.   Ti:i.iii;,'s   ^lM.\\ill^    llii'   iinns  lui    Ilir   li.ii  1 1  lic:il    ..I    Uir    Irni,'    rcMillin^    froni 
>liiiiiiliilii.n    iif    111!'    ■.yinp:illi.  lii-    iirrvi-i    prior    In    lli.'ir    iiMiiPii    HitW    tin- 

\:iHii-<    nerve 22S 

7l'.    KiU  's  kidllev  fillidllielir o;>y 

7.:.    Kiill  ct'  l.liHi.i   |.ii-ssine   rn.in  es.il;il  i.Mi  nT  tile  pli'iiri^.-...!    ],i.,\r j:;(| 

71.   The  ilVcet   el'  striin;,'  sliirmlatiiiii    (lieiit)   ef  llie  .'ikiii   ..t'  Ilie   t'not   en   tie-    ;ir 

leriiil    MiiimI    pressure    .'iml    re<pirii!ery    iMn\  iin.nl  ■. 'J  1 1 

7.'.  J)i!ij,'r:nii   shewing   \Uv    preliaMe   .•nr,Mii;;einenls   ef   llie    \a-Miiiet,.r    retli'Nes     .   LMJ 

"''•  Aertir    III I    pri'SMire.   sliewiiit;    the   ell'eet    of    pii<tiire i;4,-, 

77.  Traiiri-  u>   -liew    ih-   ilTeet    ef  },'ra\ilv    en    lle^   arterial    Mue.l    pressure      .      .  215 
7>.  Tlir   rlVeri    el'  ^.'ravity    en   tlio   aeriie    pressure   al'ler   .li\  i<ien    el'   thi'    spinal 

I'ir.l  ill  ilii'  upper  ilersal  reyieii oj,; 

7!i.  Sell,  Ilia  te  shew  the  relatiniis  el'  the   I'ai  eliieiiiau  IkkHos  to  the  sinuses     .     .  2tS 
Ml.  Tra.iiiu'  she«iiifr   slmultaneeus    n  cur.Is   ef    the   arterial   bloe.l    iiressnre.    the 
\i.ii,.iiv    pie-*nie.    llie    iiiti  11  iMiiial     pressure.    tli.>    pressure    in    the    \iiinus 

'^'■'"'^''- 'J.-.I 

s|.    i:i..lri.iar.lii.f,'iaplii.     :i|  pa ;  a1  ii-    a~    ma. I.-    Ii>     tir    < 'ainl.ri.l-.'    S.ientilie    Ma- 
terials  Ce .'i;,! 

sj.    Neimal    elerll.ii  .al.liumalii ..,;) 

>:i.   Klei-treearilie;;raiii   I'le^.  l.ikiii  viiieiiianenu^iy  wiih  run  ,  s  riuui  aiiriele  aiel 

M'lllriele .>i;.i 

'•1.    Ilei-erds   ef  iN.-l  nMar.|ie::raiu    aiel    iiie\  e nt    nf   veiiliiel,.   nf    fruL;  sheu  in;; 

that    ulii'ii   the  apex   is   aaneeil  a   lypi.al   'J'  .lave  appears    In    plaee  ef  a 
»a\e  in   the  eppnsile  , lire. -111111   app.'ariuL'  "lien   llie  ajirx    is  .i.eleil      .      .  L'i'.l 

s.',.  Sinus    lirailyear.li.a .^CiT 

*>!'..   AuriiMilar  extra-ystele _         o,;- 

•^7.    Ventriiiilar    extrasy^ti.les    arisin.;    in    llie    riylit    veiil  riile L'(i7 

>>'<.  ■\'ontrieiilar   extrasystele    arising    in    the    left    ventri.-l,.     . i.'(i7 

■>!!.   Parnxysinal   taehyiarilia Ijlji^ 

!iO.   .\nrieiilar     filirillatien of,,S 

i'l.    Auriiular    flutter 270 

il-'.  Delayed    cnnductien n-o 

'■''■'.    I'artial    dissiieiatiun n--y 

!•!.   t'einplete     dis-^m-iat  imi o;| 

I'.'i.    riilyspli\;;iii.iyrapli 271 

!lil.    Xeriiial    .iii^'ular    ir.a.in;; .j^.j 

'.'7    TJednreil  IraeiuKs  fn.ui  .areliii.  anrla.  ■.  .nt  rule,  aiiriile  ami  jUKular,  to  slirnv 

the   general    relatiensliips   ef   llie    varluiis    waves 'JT."! 

'.e^.   relyspl!y^;nio;.'ianis     imludiiii;    ju^rul.ir.    appx     ami     radial    traeiiins     ....  I'T.-i 

:i:i.   Deta.H'il    e..ii.lu.-ti.ui    time 'J77 

I'll.   Dreppe.l    beats  077 

IiM.   rrrmatiire   beats    (extrasystele  ,1    ventrieular    in   eriirin 27R 


''^^"l  ll.l.l  STIi\llll\S 

vx,.y 

liiJ.    I'ariixvsiiiiil    l.-ic  hv,  :iiili.i  oyj^ 

III'..   Aiiriiiiljir    Hiilii  r o-,, 

III).  Aiiriiiil.'ii     lliilli'i'  „-(, 

111.',    .\uri.uliir    lilirill;itiMii .,^f, 

K"'.   sl,„ui,i;r    th,.   :i|,|.,:,i:in.r    ui    111,-    M I    vr»s,.l..<    in    ll.r    r.-iis   ,,f   ;,    nil>)>it    in 

!l    -il;!!.'   l.f   ,||.,.|,    .|„„-k.        iCcil"!     I'i;ilr.  I o,,|, 

I'i7.    l>iMj;r:iiii   >tiiittiii;;  :iniMiint-i  of  iiir  .-..ntninnl   l,y   llic   liin;;s   in   viiriMiiN    j.liaisrs 

111'    iiriliiijiiy    ;in.!    iii    I'liri'i'd    ris|iir;iliiii, _  ;;(|| 

Ii'N.    l']icMin.ii:iii|,|i .,||.. 

Iiiii.    Ilcly    |,|,lliyMri(,;rni,,||    |,,,.    ri-,„r,lin-    rf«|.iiati.in :UU 

llH     i:ir,,i    ,,f   ;ilul<,riiin.'il    an.!    r)i,M    l.t-.-atliiii;!   nn    the    imiIm'   an. I    M I    |.iT.<snn' 

III'    niari .,,.^ 

III.    I'iisi    ,|...s.-,l   M.it|.|.i.-..   si\i|,   ,i,,is.l    v.'it.'l.i.-i   an.!    ril..      Axis   ,.t'    ...taliori   slinvwi 

ill    i-aili    i-a^r -.1,; 

II.'.    I,„u,.r    l.alf   oi'   ||„.    t|,„rav    t'nim    thr    I'.ll,    ,|,„>al    ti.    Ihr    till    MM, lira,    .snii 

111. in     till'    I'rnnl ■^]^ 

II::.   Inlrniistal    ninsilrs    i,(    ."iili    aril    I'.i  li    spai-.s .-H't 

III.    llanil.iTKi.i  V  s.-lirnia   In   ilniinnMrali.   t  lu'   I'lmrt  iimal   anla;.'i.nisMi   of   internal 

:in.|    ixtirnal    intrrri.sials ;;i;i 

ll.'i.  .'<.li..nia     t.i     ilrnu.nslralr     tliat     ili,.     I'nn.li f     tl„.     intrinal     intrivar 

lilanini.iis     int.nn.stals     is     i,l,nlii-al     uilli     that     i.l'     tlii.     cxtrrnal    in- 

ti.nis.si.iiiis  intrTr.istals ...i,! 

111'..   Hia-raiii    In   slmw    th,.   .fTiil    i.l'   l,ij;li    an. I    li.u    |.„silii,ns   i,|'   il„.   ,|ia|.lira;;ni 

on    I  111-    iiistal    anijlc ;;.i.i 

117.  Dia-ram    ti.    s|,.,„     tl„.    ,.|r,.,.t    „|-   Hinii-al    .lispja.i. mints    i.l'    il,,.    ,liai.lira-ni 

I. II       til.,      .i.stal      lUliiU' •jo;. 

II-    DiaL'iani  In  shu,    .uls  n-iinircil   t'l.r   isi.latii.n   i,t'  lni.   iilircni.-   .inliT     .     .     .  .'l^N 
ll:i.    Iiia;,'iaiii     li.    sIh.h     iritain     [.iisitii.ns     in     Ihr    inc.Iiilla    !iii.i     ii|.|n.|-    i-rrx  iral 
ll.l.l.    «h.'i|.    s...tiiins    may    I.,,    maili.    Hiilidiit     siTJoiisly    .jistiirliin;;    ill.' 

I'rspiral  inns ;>-K| 

IJii.    Hiauraiii     tn    slmu     HJirn.    i-iits    an.     iiiinl..    ti.    isnlali"    Ilir    rhirl     ii.s|.iiat..ry 

..  nlir    I'liini    alViri'nl     inipulsi's .•;;;n 

l-l.    l>ia;;lalM    sl,„Hii,^    |,iii„.i|.|,.    f,,,    iiirasimMiiilit    ul'    111.,    feiisi f   CO,    in    l.ji.u.l  .l.-.v 

IL'L'.   'I'll.'  ;;as  analysis  iiiprtli.   t'l.r   tli|.   miiri.'i.niiiui.tiT  sliinvn   in    i-'i;;.    IJ.",      .      .      .  :',:;.| 

\-.:.    Mi.ii.ti.iiiiiii.tii-,   I.,   1...    in-Ill. ..I    inli.   a    l.li.iiil    \  (.ssrl :i:;0 

l-l.   .\|'i.aialiis  liir  ii.ll,.,tii.n  i.f  a  samiilr  i.f  a!\iolar  air  liy   llalilam's  iiii'llii..|  :;ift 

l-.-i.    KM.|i.ri..ia 's  apjiaratiis   l'..r   m<.asiiiin-    tlii'  ('()_   in   alvnilai    air :;  ( 1 

ll-'i'..   Ciiivrs   ti.   slii.M    th,.   rclatiimslii|i   U'tHrrn    Hi,.  ()    ami   <■()_   l.-nsimis   {n    alM'i.lar 

nil    anil  aitiiial   I.1..ihI ..,1 

I:::,  .-.an,-    as     I'i;;.     ij.;.    ,.s,v|,t    ihat     i„    this    .-as,,    tli,.    triisiun    ..f    CO,    in    tli,. 

alM',.lar    air    \>;is    ,\|i,i  inn-ntaliy    .'ill,.ri'.| :;|_> 

I--.    .\ir, 111-,. Ill,  lit    „t'   I,,,. I,. IS   an,!   ,i.nn,.,-ti„ns   ,,t'    |>,.ar,-,.'s    iiu'tli.,,!    t'l.r    mrasi,,,- 

nun!   Ml   CO    ,ir'  alvn.lar  air  in  ni.iinal  .siil.j,.,ts :;i,j 

IL".|.   I'niM.    sh..uiii;;    tl„.    r,.s|.iiat.iry    i,.s|i,.ns.'    t..   CO^    in    ll„.    .1 n.l.ial,.   ,al       .      .  ::.-,l 

l.lii.   Ti'iisiims  „f  0_  ami  CO^  in  aUi'iilar  ail   at  ililVi.H.nt  altitiiili's ;;,•,! 

!!!.   Carvi-^    ■.!,„■.■.  In^;    vririnfi..ns    in    alM-uiar    cas    t.  nsi„ns    al'l,-,     r,,i,-,.,I    l,i-,.atl,- 

ini;    t,ir    lu,,    miniit.'s ..,;i 

I'.:.'.    \aii,iiis    i\|.,-   i.f    |.,.|i,„|i,.    Lic'iiliiiiL' ;.-■' 


1 


M.l.rslKMKiNS  XXIN 

►■"•■  l■^.,^: 

l::::.  <^ii:iiilil:ilhi'    n nl    iil'    liri'dlliiii;;    air    llmiii^h    .'i     ml.,'    :.'i;(i    ,ni.    loiiy    .'iii'l 

-    fill,    ill    illiiini'li'i :;;i 

l-".l.    Ifcin-rnl't 's    tiPliiiiiH'ti'i      fill     ili'li'l  liiiniho     ||,,.    .ihm.    .,1     .'il.sor  |.t  imi     of    i.wijiii 

liv    lii'iiiii^'liiliiii    or    IiIimhI lv| 

I'^'i.    Iliin-icift  ■•*    ilil1Vii-iili;il    liliiml    •.■•:.    iiiMiionii'tir :isl 

I'  >;.    liMr.ri.fl     MoimI    j;iib    iii;ii i.'tcr :;>J 

\'-'>',   Tvpii-iil  <li«viH-i;iti<'h  i-iinc.      (<*iilni-   PInti'.  ( .;s-' 

I-l"^.  .\\t'r;im'    ilisMdt-iiif  inn    rurNi's ;',^\ 

1.1'.'.    I)is<CMi;ili.ill    iliixcs    (if    lli'lnnyliiliiii US.", 

It".  Di-isdiiiitidii    iiiurs    (if    liiiiiiiin    IiIihmI ;!SCi 

I II.  (•|ii\,.s    sli.iwih^    icIiiliM'    i:il.-i    ,if    iixi.l.MtiiiM    nil. I     r.'.lii.tli.ii    i.f    I.I.1...I    :.h 

iiillui'iiicl   l.v  tciM|.i'iiitiin'  ,'111. 1   l.y   liii-.i..ii  ..f  CO .is; 

Ml'.  I'liiv.'   ..f   I'O     ti'iisi.iii    in    I.I I ;;.|o 

It:'..  I'ill>    iif    |i;ii'.itii|    ;;liiiii|    sliiiw  in;;    /.\  iini;,'iii    ;.'r:iiiiili'> Il'.l 

111.   I'ju.iti.l  yliiii.l  .if  nil. I. it    in   \;iniii^  -tiiti's  i.f  ,-i.li\ilv  .'xniiiiii.'.l   in    fi.'^li   >lnt.'    ll'.i 
1  l.">.   I)i,i;;r;iiiiiiinli.-    n'|.irs.'iil;ili(.n    nf    III.'     inn.Tviit  i..n     nf    tli.'    s;i|i\iiry    ul.in.N 

ill   thi'  .l.iy.      (<'..|..i    i'liiti'.  I (jj 

Uii.  I'iiiiircjilii'   iiciiii   slaiiicl    with    ticinatoxylin 4L'7 

117.  Till.'.'  i.;-.'|iiirntii.ii.'<  cif  i.iiiniral  ii-  a.iiii  staiiUMl  l.y  cisiiiiiran;;!.  tdlui'lin  lilui>    IL'n 
11^.    Diaiiiani    slii.Hin;;    ininiatiii'i'    sldina.li    -r|.ai;,lc.|    fn.iii    tlif    main    ^<l..niaili    l.y 

a   .I..!!!.!.'  lay.  I-  i.f   nin is   nirnil.rani' I;; I 

Il'.l.   'r\|.i.al    iiiiM'    df    snicti f    i;astiii'    jiiirc    |.|il|..|-ti..l    in    .'■  niiiiiiti'    iiil.rvais 

1.11    inastiiatidii    i.f    |.alatal'lc    fdiiil    fur   I'O   miiiiit.'S 1.17 

'■'"■  ''I'liii-    iriili liT.-.    I.f    Kfistiii-    Jiiii-c    si'i-ri'tc.j    afti'r    ilirls    of    inr.-il.    lircail. 

an. I    mil!; .(Id 

l.'il.    Di^Tslivc    |.i.H,T    of    thf    .jiii.-.'.    as    nicasinr.l    l.y    tlu'    lonmli    of    llii>    pi-otdiii 

.'oliiiiiii   .li;;i'sl.'.l    in    .Mitt's  liil.rs,   «itli   iliits  of    tli'sli.    I.ii'a.l.   an. I    milk     .    Ill 
l."iL'.   l,od|i  of   int. .slim,  nftn-  tyiii-  iiir  tlif  poitiiiiis.  iMittin^  tli.'  ihtvi's  iiiiiiiin;;  to 

till'  iiii.lilli'  |.diiion  ami  ri'tiiiiiin;;  tlio  |od|i  to  tin'  al..lomiMi   for  s i-  fiim-    111' 

l-".:i.   Till'    I'lia    u'ls    «lii,-|i    tako    plac.'    in    tin-    |M.sitii>n    of    tli.>    11.01    nf    tlir    tuiiyuo. 

Hid    sdft    pnlatp,    the    cpi^ildttis    ami     tlio    laryn.x    ilnrinrt    tlic    soioml 

s'a'ji'    HI'    sH-.'illowin;,' ^  H; 

l.'il.  N.iii  iiiatii'    onlliiii'    df    till'    sidni.'ii'li )-.. 

1."m.    Diayianis    of    .mtliiii'    .'iii.l    pusiti f    st..i h    as     ih.li.ati.l     l.y    skiaaraiiis 

t.'iki'ii    on    man    in    tl loi't    jLisiti it    iiitiTvals    afti'i    swalli.uin^;    foo,l 

iiii;in';:iiali'i|    nitli    liisiimlli    snlmitrati' .),-,■' 

l.'.l.  Onllin.'s  of  111,.  shn.l.iHs  last   by  tliv  sloniai'li  at    inti'ivals  of  an   Imni'  raili 

afliT   fi'i'ilin^  a   I'at    with    fo...!   irii|.i'<'miatoil   witli    l.ismutli   snl.niliali'     .     .    l.^vl 
1.1, .  Si'i'lidii    „f    III,,    friizcii    stdinaili    (rat)    simii'   tini,'   ;ift,'i    f.'i'.lin;;    with    foo.l 

KiM'ii    HI    thii'i'    iliir,'ii'iitly    i-oliiri'il     |iiii'liiins (.--, 

1."'^.  Miiilini's   of  shaildws    ill    aliiliimi'n   olitaini'il    liy   dximsiii'i'   to    x  rays   J    hours 

alter     ti',Mliii;j    with    foiiil    Cdiitaiiiin^i    liisimiili     snlmitrati' 4.';s 

1.".!'.  Ciirvi's    to    shdw    thi-   avi'iaKo   aK;;ri.-i!,.    U-n^tli    of   tlio    foinl    niassis    la    111,' 

small    iiiti'slin,'   at    Ihd   iIpsiKnati.l    iiil.'rxals   aftfr    tV,'.lln« l.-'.i 

111"    .\|.para1iis    for    ri'i-onlinK    ii.ntiaitions    n.'    thi-    iiitoslin,' Kit 

Hil.  DiaKr.'iinniati,'  n-prosdntatidn  df  thr  prori'ss  df  s.-Hmi'iilalion  in  111,'  inlislin,'  4.1.-. 
I"i-'.  Inti'slinal     iiinlra.l  ions    after    exiisioii     of    Hn.     ali.lominal     -.nnslia     ami 

soi'lidn    of    lioih    vny-i 41',,; 


I 


.:i!"l 


N^v-  II  I.I  >rn  \  I  iiis> 

''■'■"'■   '''i"-     '11'..  I     i.r    ix.  il:ii;..ri     ..r     I...1I1     splMj,,  Imi,.     II, Ts,,     ,,|i     til..     iiit.'^tiii:ii 

..illtl.-l.ll (,;- 

''•'•    '111'     ilt'..t      I.r     >l  iii.iiJMl  ii.ii     i.f     ri;;lit      \;i.^ii,      mi  \ th.'      iiiti'tl  Im:iI 

.■.mini' til. Ill ^1;^ 

li'.:..    |li:i-i:irii    ..I'    tii.i.-     it     l,-iK.-<     In-    .m    .;i|.-.uI.     .  nii|;iii,iin.     lnMiiiitli    t"     n.ii.li     llii. 

viiric.iis    |,:irls   i.l'    lli,.    \:,n^,-    ihti..iiii,' ,|i;p 

I'lil.    I'i;i;;r;irii    i.T    iii.'tli...|     r,,r    r.-.-..r.liii-     »t..iii;i.  li    iii,i\ .  iii,.|iis .172 

I'.r.  Ti-.-i.iii^  (iT  til,'  t..iiu>  iliylliiii  III'  til.'  vl..iii.-i.-li  tlir.'i'  In. 111-  ;itl.  1  :i  iii..'il  .  .  17.; 
Ii'."<.   'I'tih  ihj;s   t'ri.iii   til.'  «t..iiiri.|i  .liiiiii-    il iiliiiiii;it  i..ii   ..f  .-i    |.,rii..l   ..f   \  i:,'..i..ii. 

;;;i«i|i,.    liiiii^,!-    ('..iil  1:11  t  i..iis i;;; 

'''■'■   ■'^l"'»i'iH    !iii(;i"''»':ili I'    111"    kh,'.' j.rli    .liiriii-    til.'    111:11k.'. I    liuii.^.'i'    ...11 

'  i.'t.'f  i..i.^ 17- 

irn,   lii.'i.^iiiiii     1. 1'     111,,     iiriiiit'.'ri.iis     tiil'iil.~,     ilii'     ;irti'ii.  •;,     .■iii'l     tl..'     \  .'i ( 

llic    ki'liii'V "lis 

171.   I'r..^.;   ^.','ll,.n   i.r  .  ..iiv..liil.'.|    tiil.iili'-i    I'ri.m    ki.liii'V    i.j'   r.'it .".n!! 

17-.    l>in^'i;iiii     ..f    l.li.o.l    Mi|.|.ly    i.i'     M:il|,ij;ln;iii      ,'..r|.ii».l.'     :in.l     ..f    .  ..n\  ..liil.'.l 

tiiliulfs    in    !iiii|.liil.i;in    ki.liii'y ,-, |-| 

17.:.   X.'n.'    Mi|.|.l_v   of   til,'    ki.lii.'V .-.Jii 

17t.   i;,-|.ii;ili..ii     .•iili.riiii.t.'r    of     111,'     l!ii>^,.ll      Sm'^,.      liiMitut.'      ..f      l';illiol„;;v, 

i;.il.'Mii.  ii..s|'it;ii.  N.'w   V'.ik .-,:;r. 

17'".  i'li:iri    for   .1.1  .iiiiin ill;;  surfn,  .'   ;ii-.';i  of  111:111    in    >.|ii:ii'o   ho'Iits   from    w.'iLilit 

in    kiii.;,'r;iiiis   :ni.l    loifjlit     in    .I'lit  iiiiit.r:^    :ii'.'orilint;    t..    foiniiil.-i      .      .     .  .'.id 

171;.    |ii:i;,M';iiii     of     .\l\\:it.'r  H.'nnli.t     n'.s|.ir;itioii     i':)!..:  iiio'l.T ,-,.1,; 

177.    N..>,' .'li|i,   ;;i,',    i'i;i<k.  iiii.j  Mioiitli|.i.'<'<' -,7,.-, 

I7>>.    |liiii;r;ini    of    r.".|.ii:it,.n     vilhrs 3,-JC 

17!i.  Th.'    Ti.-sot    .spiioiiii'tiT r,7i7 

1^".  'I'll.'    DougliLs    I.rifj   111CII1...I    for    .li'liriiiiiiiii;;    tli,.    r.'s|,ir:it..iy    .  \.'li:iiiir,.      .     .  ^ns 

1'<1.   Ilnl.l.'iii.'   «•■'"'   ap|.!ir;itiis    :in.l    I'lnr,'.'   sriiiipliiitj   Inlo' r,','.) 

I»L'  t'lirv,.   ronstrii.t.'.l    from   .|:it;i   ol.t:iiii<.l    fr.>iii   :i    mi'ii    nln.    f;isl,'.l    f..r    lliirtv- 

oll.'      ll.nVS r„;- 

1^;.   I'iuv.'s  of  oroi'lli  of  rut.-  on  l.iiMil  r:iti..ns  |.|ii,  thi'  \:irioii>  |,r..t.'iiis  in.li.iil.'.l  -.7.; 

1">I.   I'tirv.'s  of  (,'r..wtli  of  r:ils  ..n   l.:i-.:il   r;iti,.ii<  plus  tli.'   pr..l,'iiis    iioli,:!!,,!      .     .  ,'-,77 

I-'."..    l'lMito;,'rapli«    of    r;it<    ..f    -:iin..    I.r 1    on     \;irioiis    .li..ts ,-,71) 

l>>l'.    \'l\i'iilT,i>i..ii     ;ip|':ir:it!i~    of     .1.    .1.     ,\l..'l i;ii7 

1^7.   I  iirv,,.;   slio»iii:4   til,'   :iiiioiint    of   niniii..    iiitroucii    tiik.  11    up    l.y    .litf..  .nr    tis 

-  ..'-  :.li.r  till'  ,uti.ii,'.iiis  inji'itioh  ..f  niiiino  :i.'i,ls i;iis 

l^v.  CiiM.^    -Iiowiii-    til,.    .•.ii..','iiir:iii..n    ..I'    niiiiii..  :i.  i.l    iiiii,.L;i'ii    in    ih.'    1,1 |    ,|ii, 

111;;   l:i~tiiiL.'  :iii.l  pi.it,  in  .liu,'«ti..n i;ii.i 

IM'.   l•!ll^.■s    -li'i.'.ii,:;    t!i.'    p.  1 ,-.  nln^,'    .,f    ;;|ii. •..-..•    in    M.....!    :,ft,'i    ;i    ...iKiniit    in.jc' 

ii..n    ..f  :iii    Iv    p.r  .■,  lit    M.liili..i;    ii.i,,  n    .n,'-,'nl,'ri.'    ^.in 11,-js 

'■'"•    .\ii:iii;,'.-iiM'iii    i,(    ;ippnr'itn<    for    r.  .  ..r.liiiL'    .  ..i.l  r:i.t  i..ii~    ..f    n    nli-rin.'    strip. 

iiii.-liinil  stiip,  ..I    lin^r.  i'|.' ;pi 

I'.O.   Tr:i'  in;;    sliouii.:;    Il iV.-.-l    ot    .'['in.plirini'    on    tlo-    int.  ".1  innl    .-..nt  rii.'t  ions 

.•iii'l    on    tli.'    :irl,'ri:il    I. loo. I    pr.'ssur.' .      .  711 

11'-.    .Vrriinu'.in.  lit   of  :ipp:ir:it  iis   f.,r  p.'rfii-i..ii   of  tli,.   m-^-.Is  ,.i    ;,   l.niinl.'~s   fr..;;  71.' 

llt'l.   Mi.  r.'pli..t..:;r;iplis   o,    iliyr,.i.l    ;.'l:in,|    ..f   ,|,,;r 7-1 

ll'l.  f'r.'lin.    iiin.t.'.'ii    y.'.'irs   ..1,1 7.-( 

'!''"|.  •':!> '    niy\",l.'ii.:i     l„fi,r,'    mil    :ilt.r     ii.^ii ni 7,-,.-, 

l:"'..    hr:rAin.;  from   n    pli..t.i^r:ipli   ..f  -a    iii.sl:,!   >',^iti:,)   ...ti.,,,   tlii..i:.;li   lli.'    pitiii 

t:iiy   ;;Iaii.l   ..I'   :i   huiuiin    f.'tns 7,;;; 


IK..  ,,„,, 

li'7.   Tiiii'iri;,'    !.l.pHiiii;    llir    :iiti"n    i.f     pitiiiliiii    uri     tin'    iil.'ii t  im'I  I'ln^    ;iii'l 

liliMnl    iircs'<iir('    ill    (I    tliif.' 71;, 

I'.i'*.   Tiiii'iii;;   stiMHin;;   tl lOi.-tiii-tiiiu    inti.pij    ,:(    |iin]ilii] ilir    lin.ii.ln,,!,  s   ;,ri>l 

its   i-fTi'cl    ciii    liliiful    jirr'-'^urr    in    :i   spiiinl    tlnjj ;(;(i 

l;i!'.   Sliiiwiii^  till'  ii|.i"'iir:iii.T  lu'lini'  ;,n.\   ri  I  f.  1    tlic  ..n-.t   ,,f  ai'i(.iii.>;;iili.'   ■.mm|>I(>iii>  771 

-<'■'       'mih!   (.f    .■!    |iir-ciii    :iirr,l.,|    v,  ill,    {iiiniinuidy 77J 

L'lil.    .ii,i:;t:iiii     sli..uiii^'     ^iii.iii:il     •■v.iliiii T     urn. ,11-     -\,|..|ii      in     n|„,ji-,.,     v^:, 

;(iii-irHMii'.    :iiiil    t'lirthwu)  in 7s; 

-iij.    Hiiinriiiii    (if    iiiTMMis    Mf.ti'111    111'    si-uiri.'rLlr.l     ill  \  II  t  ■  Li  a  I '•.    Mi|ir:iiMi|.li;i;;i:il 

minyli'iii.    -iilii'«ii|ili;i^i':il    ^aiiyliini,   |i^ii|.li;miis   ,,!■   ;;iil|i't 7»| 

-h:;    Si-lhjii:i    (if    -tiiiiiili.    r.'llix    arc 7s, 

-III.  TlM-iiiicii-,||i,vi,,|i,i'lir 7(1] 

:;n.-,.  r,,|i|  s|.iiN  iin.j   hi'^i    -|„iN  I.;    .,11   :iria  „(  >Mii   ,if  tif   ri;;lii    li^iml  ...  7H1' 

-i"5.    Iliii;,'i;iin    til   >li(irt    .•iMiii    rill.v    ul'   mik.hv    ihim.    lili.r   ,,r    ,kiii 7!ir 

l'ii7.   .\iliiiri/.M!i..ii   (it'   ii.llatiTMU    iinr,    1lii.    |.,i*t.Mi.r    cmi    lili -r,   nniinul    llic    c.  IN 

ill"    the    I'd-itcridr    Iiiiiii _  ^1,^ 

-'"^-    •^'"' I    "'II    f'l' Hi''    :iiii.iii.r    li.ii.i.   villi,,,., I    to    >li.ivs     Nisei's    -1:111. lie-;      .      .  7ii!i 

-'"■.'.    r.-iii    dl'    Mil    iilitiiidr   cdiiiii:!]    ell    |-,di„    ilid    ,;ilt"i    s|iiii:il    .■..nl,    stiiiin.l    In 

.-lidu     iirurdHlirils ^,1,,, 

-111.    l.i\iii:;    iiriw    iills    cviiiniid-.l    l.y    lln-    nil  i:Miii.Tdv.d|i.. siij 

-'"•   '■''•"■II'-    I'li'iM    Hid    l.iii.j    liinl,    i.r    ;i    ,|,iii.-i|     1,1^    ilniiim    tin.    ,-,-iiil,l,in-    1,11, V,. 

iiniils    iiniiiiiiTil    l.y    :i|i|il  v  inu'    »lii.iiili    ;it    Ih..    -kin    imiiilv >.|J 

I'll'.    l,',r,,i.l    t'lHiii    iiiyd-i:,|ili   , .,.i,.,|    uiih   tlid   cNliiisdr   innscld   dl'   Hid    knee  ^l.•| 

-I':.    Iliimr;,„i    vliduin^'    tl.d    miisild,    nn.l    ii.rvds    .•diir,,  ,id,|    i„    riri|,,,,i!,l    iniici- 

Mllidll ^|,; 

-II.    Ii'i  ii|iliil;il     illlliiv;il  iiili        ....  S|- 

-I.-|.   Shiri-iii'^ldn's  ilinyinni    illii>trni  in.;   ilir   inr.|i;iniMn   nl"   rc.i|ird,':il    iniidiMil  ion  ».s 

I'Hi.   Ili:i-i:iin    slmu  ir:-    il,,.    ivllrx   n  n  -    ih.iilvi.j    in    il„.   s,i-.-ii  ,.|i    rdflrx      ....  v.'J 

-'17.   .-ihinvin;;  rv-ii.,1  ,,1'  ho.ly  „f  ,l(i;r  ,■,,,„,   „l,i,li  tlir  .rriitil,   ivIIia  .nn   1 li.-itcl  v':; 

-I'*.    I>i;i-ii,iii    >li(i«iM-    tlid    scjini.nlnl    mi  i.-m.^diiidi.t    ..(    llid    s,„.„,v    ncrvdv      .      .  s:;7 

-'III.  Oiitci-    ;is|i,.l     iiT    thd    liriiin    nl     llir    Hiiin|.,-iii/dd      ....'. s(7 

L'l'ii.  'Miiv,.   ■idciidii,  ;lin.ii-li    .lilT.r,  11'    \,:nu  ni    ilir  ,rr,l.r:il   cutdx ,s.-..' 

l"-'l.   Till-   Idiiiiidii  df  111,.  ,.|,i,.i'  ,„,„,,,.  .|„,i   „.n,„,y   ;iii-;is  dii   111,,  diitcr   ...n.l    nu'^iiil 

Ms|.d(t-i    df    !lii.     liiiiiniii     l.iMiii S-. 

l|Ji'.    |-ddt|„iiii.    :,ft.i-    .|,.,i,ii,ti„n    ,.{■    111,.    ,-,.,,.l„.ll,nn    in    ;i    ilo^'     ......  si;,; 

-'-■'..    hi.'i^inn;.    Id    IV, „,..,. Ill    i(.,,,i.,iiv,|y   „    x,.,„,mI    vi,u    ,.f   ll„.    l,f,    l,,,!,'   ..,,,,1   n 

.ldi..-,l   vi,u    df  III..   .i;,l,,    i,„||'   „,•   ,,„,   ,,,„„.,„   ,.,,r,.l„.i|,„„   illiistr.-itin-   tlic 

-.  Iii'iiid    df    siili.li\  i-id,i    111  i-drilinL:    t.i    lidli; sil> 

-L'l.   .S-ldinn  df  ,|„.  |,n-N  ,.f  tlir  i„.-n,i,n.-,li:,n  (■.■ivlnlli,,,,  spivml  diit    in   dnc  |.|„nc  '  Ml'l 

--"   '■""'   --'•'   ■'''"•    """'"l^'timl   iinil   111,.   iiiiMnidi-   ;,., ,    ,„■   ,|„.   l,„n,;„ ,„,„,„„„ 

""'"■■'""-    ''itMin    .•,Mdli,.ll:n     I,      li/„ii.,n.i    ; ,,li„^    ,„    Hnr,-ii.v     .     .     .  s7i 

--'■   ■'■'"■    -''"'i-i'iiil'"'    '"n^U    „f    t! ,.,r.    sl„iu  in^    tl„.i,.    :,,-r:,nf;dn.|.n't     in     t!,.' 

till'..'    iil;iiid<    df    ^.|,:,|.,. _  ^.| 

-'1'^.    ni;,mnin    illnsliMtinu'   tin'   ilifl-cMMit   iiiruiu.iucils   .,1    tln^    int,.n,;ii„.'i„l'  m^iimn^ 

dl    ;lii'    Milriilniy    mil!    iiiMiluiii;i,y    nrrM.us    sysicins S7<! 

--II.     hiniilail,     i,f    llid    sv. ,,;,:, 111..!;,,     „rr^,:■:■i     ,;,;.•, »     »..    |.,.     !.„.,]     .,;,„,..     ..:,:       ],•;.'     .'....' 

■'•dldi     I'lnlc,  ,     .    .  ■■  •    ■    '~    "d"       I-.   _  •-. 

X  ,  ^ 

-•"■"■   "':'«i''"ii    '■li'iNvin-   the    mm r    i.f   .■diin,,.!  imi    „:'    |1„.    lili,.,-.   ,.,„„|,„.i„j;    ,|,„ 

iridllt     -i|,l;i,|r|nii.-     IIIMVd.         iCmIii,      I'!;,,,,     , "  ^.^ 


li 


Il.l.i  siij  \  I  hi\- 


'J-''\.  hiiiulMin  »liiiH  iiii:  111.'  111:1111111  in  whiili  a  |.i  .-iiii^linti  i.-  lilM-r,  riii;iil!lt  ilitf 
liiiMi  llii'  -iniKil  riiiM  |.\  III,'  whiti'  1:1111111  I  iMiiiiiiiiiii'iinx,  1  iiiiiii'i-N  in 
:i  i{ani;lii.ii  nt  tlir  -\  hi|.iil  In  i  i,-  .  IkiIm  i\illi  ;i  11.  1  \  ,■  ..11,  llic  nxi.ii  ..f 
v\  lii.li  III.  II  |.r...'.  I'.U  n.  III.'  |M.«l;;:iii;;li..iii,  tili.i  In  » m  ,,t'  lln'  tif.iy 
r.iin.i«    .  .>iiiiiiiimI.':iih    l.:k.  k    t..    lli.'    i{.iii:il    n.n.',    :il.inL.    wlii.ji    il    tr;u.'l-. 

I..     IIm'     |H'ri|'l|.    I  >  .  ll'li|..1       rilltl'.   i SMI 

-'^!l'.    Iiiiiv'rnin    >.|i.iv\iii^    III.'    iiiniii    (..-nt^   i.T    lli.'    :iiitiiii..inj.     ii.'i  \  .nis   s\~li'iii    l.i    !..■ 

11-.'. I  iili.n-   uitli    I'lu.  --"'.        I'lili'i    ri:il.'  I svL 

l':!:i.   S.lii-iiinti.     i.'|ii.'.i'nliiti..ii   ..1    111.'    inv.'liintiiM    inrjciis   .v.lini.       iC.l.ii    I'hili'.  1    ss  I 


rn4YsioLOGY  AND  biochi:mistry 

IN  MODERN  MEDICINK 

I'AKT  I 

TIIK  niVSI('()(  IIKMICAI.  lasiS  OK  I'llVSlOl.OdlCAI. 

IMiOCKSSKS 


<  ii.\iTi;i,'  I 
<;knkk.\l  (  (».\siih:i{.\ti<i.\s 

Tin-  witiU  iif  the  |ili\>iii|nyist  i >isis,  in  |jii'j:i'  luiil,  in  ;iM'crtiiiniii;r  to 

ulijit  cxlcnl   till'  Kiiouii   \:\\\s  i<(  pliysirs  mmiI   cliciiiistiy  liih'   ii|)pliciili()ii 

in  cxpliiininir  the   |ilii  iionitMiii   nl'  litV.     Me   i:;iliiiTs   I'r tlic   v.ist    store- 

lionsi-  of  pliysiciil  ;iii<l  ilii'niic.il    know  IimIuv   w  IuiIcmt  is  of  viiliu-  in  tin' 

inli'ipn-tiition   of  ilic   Mirions   mm clniMivnis   tiuit    woik   toi^rtlnM-   1o   < i- 

liosc  till'  livinir  iM.ii-iiinc.  tind  li;i\in'_'  inlili'.l  to  tliis  know  Itilyc  In'  passes 

it  on   for  use  Ip.v  tliose  wlio  are  e ineil  in  l|je  stinly  ami  treatment  of 

<lisease. 

Many    of    the    most     important    steps    in    the    aiKanee    of    pliysioloi;ie 

knouledu''   in    r nt    \ears  have  ilepended    npon    the  diseovery   of  sonw 

hitherto  nnknoun  ph>sieal  or  eheniieal  law.  oi'  upon  the  eialioration  of 
sonje  aeeiirate  mi'thod  for  the  measni-eiiient  of  the  )>henoinena  ill)on 
which  these  or  previously  known  laws  depend.  The  dis<'overifs  of 
\an''  Iloir,  Arrhi'nius,  and  Ostwald  of  the  so  called  laws  of  solution 
were  ^.M,n  followed  liy  iiriportaiit  ohservalions  on  their  relationship  fo 
the  movem.'nt  of  llni.ls  and  dissolved  snlislanees  throuudi  cell  mein- 
liranes:  the  diseoveiies  of  llard\,   Willanl  (iililis,  etc.,  of  tin'  liehavior  of 

colloids  and   id'  the   phenomena    of  surf; tension    found    application   in 

cNplaiuiii','  many  hitherto  ine\pliealp|e  peenliarities  in  the  activities  of 
ferments;  the  disc,, very  In  Xernst.  etc,  ,,f  melhuds  f,>r  tlie  meaMiiement 
of  the   ele.-tro motive    foice   of  dissolved    snhstan.-es    was   applii'd    to   de- 

teimine    the    a<'tual    reaction    m-    h\  droiren  ion    i ceidr-ation    of   animal 

1 


I 


Il\>n  1)1  tii'Mii  \i.  |l\sl.■^  i>y   I'llVMiil  iHiii  \i    |'|iii('i:smi:k 


Itlliils,  atiil  to  r\|i|||ill  tl;.'  Ki'lirl'iltiiMl  iiC  till'  cli'i'l  lie  riiri't'llls  uliifll  lie 
<'iiiii|itill>    Itill^i'iiliii'.   iH'I'MillH,   iiikI   ulMlnlilliil-  iirlivity. 

It    Wdlllcl    III'    lillt    til'    pllirr    IllTl'    til    lil'Mitc    IIIIH'll    '«|>lll't'    to   It    tll'tllill'il    111* 

ciiiiiit  III'  siii'li  in.'ittcis.  Tlicy  lii'liiiii;  iiinic  |iiii|>ril\  in  tlic  ijiiiiiiiiii  nl' 
Ui'iiri'iil  tliiiii  III  tliiit  III'  liliiii.'iii  |ili\  siiiliiuy.  <  ii'iii'i'iil  |ili\Miiiliiiry  Ih  i'iiii 
I'lriii'ij  with  till'  stii'l\  111'  till'  rsM'iitiiil  iiiiliiii'  "f  tin'  vital  piin'i'sscs; 
wlii'I'ras  lililiiaii  pliv  sjiiliiiry  is  iiii'ItIv  a  liraiicli  nl'  tlir  slllijri't  ill  wllil'll 
MlH'cial  attriitiiiii  i^  iliMitril  tn  till'  ap|ilirntii>ii  nl'  tin-  tnitlis  nl'  t'ciii'i'iil 
pliv  sinlniiN     tn    tlir    w  nl'k iiii;    iif    till'    lillliiail    lliarll ilir.       {''nr    till'    pliysii'lail 

ami  Mii'iri'iiii  a  Kimw  Irilirr  nl'  IniinaM  pli>  sinlnuy  is  as  issi'iitial  as  is  a 
kimw  li  .I;;i'  nl'  llii'  riiiist  nirt inn  nl'  a  pi  nl'  iiiai'liiiii'i'y  I'nr  tlir  i'ii!;in('i'r 
wlm  attriiipts   its   iipaic.   Imt   nli\iniisl.\    In  ai'i|iiiri'   this   kimw  Iriii;!'   the 

I'Minlaillrlilal  plilii'ipli'S  nl'  ui'liiial  pliv  sin|n>_'y  lllllst  lilsl  nl'  all  In-  lli:'Iir 
stnnil  I'ni  tlirsi'  ii'Msniis  till'  illt  Indllrlniy  rhapti'ls  arr  llr\ntri|  tn  a 
liiii't'   ir\  irvi    nf   till'    iiiiist    iiiipnrlaiit    III'    till'    pli,\  sii'iii'lii'iiiiral    priiii'ipli's 

llpiili    uliifh    till'   Wiil'Kili'.'  Ill'  th 11   <lrprlii|s. 

I'iniii  till'  vii'Hpnillt  nf  till'  physical  I'lirlilist  till'  nil  i-nlisists  nf  all 
rlivi'inpi'  nf  iiiiiii'  111"  Irss  pri  iiiialilr  liiatcri.'il  iiirlnsini;  ;i  iliiiili'  siiliitimi 
111"  rrystaliiiii'  siilistaiii'i's   in    which   rnllniil    inattiT    is  siispi'mlcil.      It    run 

tains,     ill    lillllT    \Mi|i|s,    a     snllltjiill     nf    i|\s     lllniils     allll    rnllniijs,     ill     wllil'll 

thrsi'  all'  in  a  stair  nf  ri|niiilii'iiilii  with  I'ai'h  nihil'.  This  ripiililiriiiiii  is 
ii'ailily  alti'iri!  y  vaiimis  inllin'iiri's  tli.'.l  ina.\  ai-t  nn  ihr  i-i'jl,  miil  tin' 
i'isiiltiii!»  I'haii'^is  iiiaiiifrst  IIu'Iiisi'Im's  niitwanlly  li>  altiTatiniis  in  Ilir 
sliapi'  ainl  \iiliiiiir  nf  Ilir  rrll  finnrtli  nml  iiinliiiii;  liy  tin-  I'xti'iisinii  nf 
sniiit'  nf  its  I'liiiti'iils      SI  in  lint: ;  nr  liy   llii'   prnpairatimi    tn  ntlii'l'   parts  nf 

till'   ri'll,   III'   its    pn SSI'S,   nf   till'   silitr  iiistllll».[    t  i|uililil'illlll       nimiiis 

liniiiilsi.       hi'siijrs    till'    artivitirs    that    aii'    ili'prllilrlit    lipnii    physii'iii-lii'lii 

iral    I'lianu'is,    piiii'l\    rhrinii-al    pf ssi's   l'o   nil    in    tlii'   I'l'll.      Many    nf 

Ihisi-  I'liiisist  in  till'  liiiakilnu  II  alnl  ii\ii|atiii|i  nf  cniiipli'v  iiiistahli'  nl'!;aiiir 
liinlri'illrs.  a  pini'iss  iilililiral  with  that  ncrliriilii;  ill  i-niiilillstinn  nutsiiji- 
the   ri'll.     Others   iiniihi'    till-   Imililiiii;    up,   sla'.'r    liy    stairi',    nf  i-nmpli'x 

slllistallri'S    nut     nf    the    I'li'llllllts    III'    mil     nf    silllpllT     Illnjl'I'llll'S.        ( 'lll'lllil-al 

tiaiisfiiriiialiniis  nri'iir  in  llii'  itII   whii'li.  in   tin-  I'lu'inical   lalimatnry,  it 

i(ilirr  llii'  iiinsi   piiwi'ifiil  rraj^iiits  ami  ph,\sii Iniiiical   I'nrrcs,  ritlii'r  tin- 

st  rnii'.'i'st    nf  aiiils,   alkalii's,  n\ii|i/iii'_'  aui'ii's.   I'tc,   nr  rxtii'inc   ili-u'roi's 

nf  hi'at,  rii'i'tl  ii-al   rlirl'sry,  I'tr.      lint    this   is   lint    all,    I'nr   ill    th II   tlll'si' 

rhciiiical  1  ran- rnriiiat inns  arc  capalilc  nf  liciip,;  ^uiilci]  tn  a  very  remark 
alilc  ilc'.'ii  c  nf  iiii'it\    sn  as  tn   piniliici'   iiiti'iiiiciliatc  )irniliicts  thai    arc 

Used  I'nr  sniiii'  spcii.il  piirpn- illici'  li>    the  cell  that  prndnccd  tliciii  nr, 

after  traiiNpnrtatinii  liy  the  lilnml.  etc.,  liy  cells  in  nthcr  parts  nf  the 
nriranisiii. 

It   is  ciistniiiar.x    1(1  spi'al<  nf  tl I!  as  a  chcinical   lalmratiiry,  Imt   it 


i.\\\w  or  »(ii.i  iiuv 


i»«   llliifl'   tlllHI    thin:   it    is   II    lillMllllliil  \     till  hi^llt'il    Milt    IIMJV    uilli    tile   ciiuiii 

MH'iif   (if  the   (lifiiiist    lint    .lii.M't.'il    ill    ill.'    Iiiiiiiiiiiii>iii>4   ii|iriati<>ii   i>t'   its 
iiiiiii\   tii'ti^  ilii's  \i\   II  ituiiliiiu  liiiiiil  vt  liii'li   till'  siirpassi's  iiii\  thini;  kimuii 

to  lllllll.     ('Ili'tllH'  'I  tl'illisl'olllllltiotis  thill    li'i|llll'i'   till'  tlli'ir  1 iililpllslllilfllt 

tlir    Jtrfiiti'sl    skill    pli.r.'cil    uitliiilll    ii|i|iiiHMt    ijitliiMllty    in    ll 11.      To 

wliiit  iiii-  llirv  rlMin.'..s  (In.'.'  What  is  tin-  iiatiiri'  «(  th.'  .'Iniiiiral  nil 
ucnis  ami  I'.nc.'s.  an. I  wluit  is  the  .jin'ctix  <■  iiiiliii'iici'  that  niii.lc's  lln-ni 
ill  tlli'ir  \aiiiil  activ  itirs  '  T..  tln'sr,  whi.'h  ar.'  aiiinti):  Ih.'  yri'Mt  ipii's 
tiiiiis  III    L'i'iiiial    |ili\si.i|i.i:\ .   till'   ii'|ilv   Mia.v    lir  ..'uni   that    tin-   ri'um'iils 

all'   111.'   iVi'iiii'iils  1.1-  iii/Mii.'s.  ami   lliat    tin    iliii'i'li\  i-   inllni-i ipccatr^ 

lliic)ii!.'li   till'  siisi'.'ptiliilitv    ,.i   (.|i/\iiiii'  arlivili.'s  t..  i-liaiiifis  in   the  nivi 
iiiiiliii'iit   ill  Mhi.'li  llii'  .'ii/Miii's  ,11.'  .ic'tiii'.'.      In  iiiaii\   ras.'s  tin's.'  .'liaiiK.s 

'■'II'    '"•    'Njiliii I    .III    a    pli.vsi.'.M'ii.'Miiial    liasis    as    il.'p.inli'iit    np.in    Hi.' 

kiniuii   hm  s  III'  mass  ai'limi  m-  siiria..,.   inisi..::  ;   In   ..tli.'r  .'asis  llii'v  i|.- 
pcml   .III    piir.'l,\    .'lii'iiiii'al   .'hati'.'.'s   m   ih,'  .'.'||   .•i.ni.'nis,  sin-li   as  chaiiirt's 
ill    irai'lii'ii    III-   the   ai'i'Miiinlatinn    <<\'   .'li.ini.'al    siil.staiii'.s    thai    art    liki- 
pi.is.iiis  .III  ih.'  ru/.\  nil'.     Mm  thiTi'  ai.'  slill  .iih,  is  that  app.ai'  t.i  ili'pcinl 

"II    iiilliKl ^   \>hi.'li   MS  yi'l    arc   .|iiil,'   iiiikmiuii    In   tin'    physi.'al    clii'iiiisl . 

Mlrh   as  till'  I'lianifi's  in   ,-,'1'   a.-iiviiv    that   ran    In'   l.r.niirlit   alMiiit    li\    th.' 
n.'r\i'  iiiipiilsc. 

Thcsi'   pivliiiiinarv    ninarks   v\il|    s.'rv.'   I.,   imii.'ali'   ih.'   pnililciiis   with 

vvhii'h   «.'  iiiiist   first   iipv    ,,iir  ali.'iili.ni       Th.'v  .•iin.-.'rii   the   pli\sii-i.- 

••h.'iiiii'al  naliiii'  .il'  saiiii.'  snjiitiiins  ami  nt  <',ill..i.|s.  ami  th.'  uciifial   iia 
till-.'  .if  <ii/.\nit'  a.-tiiiii.     Till-  kmiul.'ilu-i'  whii-h  u..  a.-ipiir.'  will  li..  I'.inn.j 

'"  '"■  "'■  ^'iliii''  ""'  ""l.^    ' ;iiix.'  il  "ill  h.'lp  us  1,1  nml.'i'staml  th.'  natiir.. 

I.I'  th.'  \.  ..i'kiiiLs  III'  ;1,.'  mirii;:,:  :, ...^liliy  .'I'll.  I, 111  iM'caiisc,  liiT.-  ami  thii.', 

it  will  imli.'ai.'  p.issiM.'  .'aiis.'s  |-,,r  .l.'ran.,'..m..iit  ii Mnlai-  |'iincti..n  ami 

Mii.".'.'st    latiiiiial   niraiis  liy   whi.'li   w,'   mav    all. nipt    I.,   ici'tifv   tin-   fault. 


} 


THE  PHYSICOCHEMICAL  LAWS  OP  SOLUTION 


The  Gas  Laws 

Thr.'.;   Mimlaiin  iital  j.rim-ipl.'s  ..f  .jcm.ral   .'h.'iiiisliy  s.'rv.'  as  the  l.asis 

I'm-  an    nmli'isiamlin-  .if   th.'   naliir '  s.ilnii.ms.      Tlii-   lirst    is  lliat    if 

Ml'  tiik"  a   .|iiaiility  ..f  aii.v    '.^as  .'.iiial   I,,  its   iM,,l.'..||lar  w.-iirlit    in   ^mmiiis 

"'•'"'■''  ''  '-liiiiii  inul.'.'iil ■  fur  sak,.  ,,r  l,r,Mi\    ,•,   n,,,!    ,   it    ^^\\\  ...'.-npy  ,'n 

iK'tly   224    lit. Is   at    staii.lar.l    t.'inp.'ratnr.'   and    prcssiir.' ;   Ih,.   s,..'.,ii.l    is 

that,   as    \\.'   .'iiiiiiir.'ss   .-i    ir:iv     its    i.i'.'ss".-,'    •''■>>    '■ ;■       v      .'      .i 

saiiii'priip,.rti,.ii  as  th,'  v.ilnni.' .liniinisii.'s  ith.'  v  ..liiiin.  „,' ■,  .as  is  invci  s.'h- 
pn.p.Mti..mil  1,1  its  pi-,.ssiir,.    ;  tin-  thir.l  is  that  ,'il|  ..ras.'s     ■  ,,aii.l  l,v  I   27:1 


I   It 


'  J 


4  lllV^lronllMIc  Al.    IIASIN    llK    I'llVNiul.lKIK  Al.    I'H()<  IISSKS 

piiit  III'  tlirif  Miluiiif  at  H  ( '.  lor  i'\t'ry  ilcLrrt'c  ('.  lliat  llicir  tt'iiiperatiii't' 
is  taiscil* 

Till'  iircNsinf  of  a  uas  is  rinMMiiiil  l)y  loiiiiccliiii;  a  pi-cssiirc  }jaiii.'<'  or 
inaiiiiiiii'tt'r  with  tlii'  xr^scl  « liidi  ccmtaiiis  the  uas.  Nnw.  it  is  plain 
that  it'  till-  ■_'■_'. 4  liters,  wliicli  is  the  Mijuiiif  iiiMMi|iic(l  l)y  a  irrain-iiinlccnlar 
i|iiaiil  il.\ ,  ■.\  rrc  I'Dinprfssc'cl  sii  as  til  iicciipv  a  viilimic  ol'  1  liter,  its  prt'ssiiii' 
w.itiM  111'  ■_'■_'(  tiiiii's  lliat  111'  1  atiiiiisplii'ri'.  oi-  ■J2.4  TtKt  miii.  Ili;  the 
ti'iiiperal  lire  reniaiiiiii'j:  eniiNiaiil.  t'luler  these  ediiilitinns  we  iiiiist  iin- 
airiiie  that  the  iiiiilei'iiles  iil'  'jas  are  crnwiled  tiiiri'tlicr  li\'  the  ciinipressiori, 

ami   it"   \v  e    I'lirthei In-eive   nl'   these    iilnlei'llles   as   l)eili<r    ill    eiillstailt    )no- 

tidii.  then  \\  e  call  iili(lerstalii|  w  h v  the  pressure  shnllhl  iiierease  just  in 
pi'iipdltimi   as  we   eiiiiline   the  spaee   ill   «  liieli    they   ejm   miive. 

(•lie  iitlier  |)ri;perty  111  uases  iiiiisl  lie  liiinie  in  iiiiiiil  namely,  their 
tcliileiiey  tn  ililVllse  rrnni  jilaees  where  the  pressure  is  hi'.ih  t(i  plaees 
where   it    is  lii«    until   the  lu-essure   is  the  same   t  hniUirhnut. 


OSMOTIC  PRESSURE 

These  ruiiilaiiienlal  I'aet-N  ri".^ari|iiie;  the  liehaviiir  of  uases  siiee-esteil 
to  \airt  lliilT  the  hypiilhesis  thai  muh  nih  ^  nf  difs^ol  i-i  il  Kiihstiiiiri  s  iinist 
Ixlmri  III  It  siiinliir  iKininir  In  tlin.--i  nt'  leases,  Tii  |iiit  this  hy|i(ithi'sis  tn 
the  le>t.  it  is  neeessarv  that  we  ha\e  smiie  mcthml  I'nr  iiieasiiriii'jr  the 
pressure  111'  ilissiiheil  mnleeliles.  W'e  eall  lint,  as  ill  the  ease  nt'  a  i-'as. 
Use  an  nrdinary  maniniietei',  I'nr  this  wiuihl  measure  mily  the  pressure 
III'  the  siihent  nil  the  walls  nl'  its  enlitailier  allil  Wnlllil  tell  Us  imthilli;  nl' 
'he  |iressllle  nl'  the  ilissnKeil  mnleeliles.  We  must  Use  snlile  (liter  nr 
llieliiliraiie  that  will  allnw  the  llinleeules  nl'  the  snivellt  lnit  lint  thnse  nl' 
the  ilissiiheil  sulistaiiee  tn  pass  thrnll'^li  it.  It  is  eviilelit  lliat  it'  siieh  a 
tiller  is  iilaeeil,  I'nr  I'Nample.  lietwei'M  a  snlutinii  nt'  suuar  ill  water  anil 
water  alnlle.  the  mnleeiiles  nl'  the  latter  will  (lilTllse  illtii  the  snlutinii 
until  this  lias  liei-iime  sn  illlllteil  that  the  pressure  nl'  till'  llissnlv  ed  mnl- 
eeliles is  eipial  nil  linth  sides  nl'  the  inemliraiie.  Siieh  a  liiemliraiie  is 
ealled  SI  iiiiji,  i-)iii(ihli  :  the  dilTiisinii  111'  llinleeules  thrnUeh  it  is  railed 
iisiitiisis.  and  the  pressure  whii-h  is  eeiieraled.  flu  nsiiinlii-  jin  ssiiri .  1 1' 
we     pli'Velit     the     water     lllnlerules     rriilll     aellially     dilTllsillU'     li>      nppnsine 

a  pressure  w  hieli  is  eipial  tn  that   with  w  liieli  they  tend  tn  ditTiise  thrniiirh 
the  liieliiliraiie,  w  •'  eall  tell  the  lnaynit  llile  nl'  the  nsinntie  pressure  I  l-'i<r.   1 
III  applyiliii  these  I'aels  In  test  the  hypnthesis  that  mnh'eilles  in  snlutinii 

•Tills  in-ill,-  Ih.it  -,l  .';■."  ('.  ihv  i:-,-  «..il,|  ..,  .ui-v  iif  v.. Inn.-.  Ililni.'  Ih;-  Iiiiii  ii.itnr.-  !- 
na,  licl.   lii.ui'v.r.   llic  li.|nrf.ul|..|i   ..l   Ui,    1:^1-  -,  t>   in.      Tin-  li-iui.i  i.ilnrr     .'.".i"    l'     1-   l>i.n«n   av  .il.-,  ,lnl.' 

'•■■■'■■     -Ar.   : '■      ■■.,.:  I. :.,[ ,  i.iii.r.   /,■,.,    ■, ,;     ,-  ..,;■„,',  lii.    .,;  ..,;,,!.    i.i(,|..i.. ...i;!.  1   «.n   ,11'  »i.,i 

uii:  I'll-  .,i..,\,     Li,>    ■-   tl-.-Tcl,.i,-   ilMi    ;!•.    ^,,lunl,     1-   .\t  .-tl-.    .......,.v i.il   i..   ll,  -..'ul.    l.-Tn|.,  r.iliir. 

A(   ;7.r    l".    thr    M.liiinr   nf   a    K.i-   at    n      C     W..11I.I    l.«-   .lniil,l,-,|,    .,r    if    .vi.an-.  ri-    i.r.i,  iit,-.l    111. 
fir. -Mirr  «.'ii!.l  ll.    ili.^iM,  .1 


I.WNS   III'   SCl.ri  KIN  •> 

nlicy  the  sii'iic  l;ius  jjs  IIimm'  in  irjiscuiis  Idrni.  \vi'  must  cnililoy  ii  sciiii- 
iMiiiii'.ililf  iiH'iiiliiMiic  \\liicli  is  liiriil  iiiniiuli  111  w  illistiiiiil  llii'  |)i(ssmc 
ami  uliii'li  fiiiius  |ijiil  nl'  tiic  ujiJIs  n\'  ;i  i-iosril  Missel  fomn'cti'il  uitll  il 
iii.'iiiiiiiit'tci'.  It'  w  r  |il:ii'('  ill  such  an  iisiiixiiiclci'  a  siilutimi  rniilainin<r  tin' 
riKilciMiiaf  wciylit  in  yrai'i-  of  sonic  siilistaiii'c  ilissolxcil  in  cmc  litcf  (if 
siilvcnt,  a  sii-callcil  ;'r,:;i.  !.i.,U'v,i',n'  suliitinn,  it  is  oliviiius  that,  il'  the 
'j-.'is  laws  arc  tu  appl  .  liic  usiii  ilic  ;  i  ■ssiirc  slicmM  .iiiial  that  nf  '22.4 
liters  of  a  u'as  ciiiii|ii  -.  ■  li>  llic  \i  'iiic  nl'  niic  litci-;  in  otiici-  words, 
it  slniiihl  cipial  ■_'■_*. 4  •  ,  ■'  li'iii.  Mj-  Jtlniiiuli  there  are  very  ecmsiilcr- 
alile  tcehnical  ilil'llenit  ies  in  iii.il  ,:<"  a  seiiii|ieriiiealile  iiieiiiliralie  that  is 
vtiiiiiu'  ciiiini;h   tci   withstaihl   such    a    pressure,   yet    this   has   In  en   aeenni 


n 


I'l.-  I  ll|,,-Jl.ll.i        Ml         ..M.I..II;,   I,   t  I'll,         ,>l|lli|ll..ll        V,-.l         MM,        ,n|l,  I        l,.ll,„ll        ,i|         11111:1, IM, I 

1I..V-    llu      I'Mi,,    ,,t     uhi.li     an      III..!     ,,nu    a     '.i .  .  n.il..Ir     ..I     <..|.|.,i      I,  n ...  «  .uit.t.      t..     eiir     .1     ^-iiii 

I'niiMil.l.      11'.  11.1.1.111,  .     i~    -.1-1 !..|     111    a It.  I      ^.— .!,    an. I     i-    .i—.l     .i'...\.      In     a    liuhnv      lillliiK 

-I. .|. I'll     piiM.-.l    l.%     a    tii:..      l.-ijiiic    I.,    a    1 1..1...1.I     .i.'i         .1    ...main-    .>    -'■.■n,;    -..liili.ni     ..I     .anc 

-v.r.  an. I  II  ...i.iain-  u.il.i.  I  la  wal.i  111..I.  .  ii!.  -  I.  11  S  I..  |.a--  lliMimli  lli,  -,  iii.|.,  1  m.  al.l. 
'.m'.'i".  Mil..  111.-  ,ai..  -iiL'.ir  -..liili.,11.  a.i.l  -111..  ill,  ..ri.  -iljai  ill.  .1.  .  'iL  -  ..ill  ml  |..l--  111 
I'"     ..I.]..  Ml..     .Ill, ,11. .11,     111..     |.i.--.ii.      in     II     11-.-     all. I     1.     .....|.l,.l     111      ;;,        ■|'lll-     ..|llal.     ill.      ..-ni..ll.- 


|ili--liei|,  and  the  riindaiiieiital   |iriMei|ile  has  thcrerurc  liecii   lirmly  estali 
ii^lnd   that   siilistani'cs  in  snliitinn  nliey   the  same   laws  as  u'.'ises. 

I'nrlher  Iinmr  that    the  i;;is  laws  ap|il\    tu  snlulinns   has  liei'ii   secured   liy 

■  liiiw  iiiij;  that  the  iismiilic  pressure  iiil'  a  dilute  soliit  inn  I  is  dirccily  prn- 
|iMi-!ioiijil  to  the  coneentrati.iii  of  the  dissolved  siilistaiice  itlic  soliitc'l 
.Hid  to  the  alisoliitc  teiiiper;it  lire.  It  also  olieys  the  law  of  p;irlial  Jircs- 
^iiies,  which  states  that  the  total  pi-.-ssiirc  exerted  liy  a  iiiixtiirc  lof  (r.'iscs 
III-  dissolved  iiiolecidcsi  is  till-  sum  of  flic  ))rcssurcs  wliich  caidi  eonstit- 
II' III  oi  the  miNtiirc  woiild  exert  were  it  alone  pi'csent  in  the  space 
'"■iiipied  Iiy  the  mixture. 


i      if 


i   i! 


G  i'ii\>-iriMiii;\iii  \i,  I'.vsis  ui'  I'ln  sKii.iHiicM.  I'UiM  i;ssi;s 

Silii'c  llic  iiMimtii-   |iri's>illr  is  ;ili;il(ijriiiis  to  llic   pifssiirc  i<(  tl  '_Ms  .'liiil 

is   tlu'ifrmi'    pi(i|icjitinii:il    to  till'   iihil'(Mil;ii ih-imiI  liititm    (i.e.,   iiimilicr 

(if  iiinlcc-iili's  ill   unit    sp: ,  it    t'lillous  tiiiit    ;i   siiMii>''iiiif.ilp|c  iiiimiiIummc 

I'.-iii   ill'  used   til  liitcniiiiic  llic   rrhitiv iic'iit  riitinii  nl'  Iwn  siiliitions  nt' 

tlic  s.-iliic  slll.staiiri'.  AVJH'ii  ;i  \\;iti'l\  siilulinli  .if  siillic  Mllistillicc  is 
placi'd  ill  Mil  iisiiKiiiii-tiT  tliiit  is  siirniiiiMlrd  liy  M  siiiiiliir  lint  iiicirc  'iiiitc 
snliitimi,  watiT  iiiolcciilrs  will  ililTusi'  into  tin-  (isnioiiiftiT  until  the  ]>ri's- 
siiiT  is  i'i|lial  oil  the  two  siilcs  ol'  tlic  scliiipi'nucalili'  llicllilil'aiir ;  that  is. 
llic  water  will  pass  lioiii  the  solution  lia\iii'/  a  lower  osiiiolie  jiressiire 
into  the  solution  liaviiiL'  the  higher  pressure.  Wlic^li  two  solutions  lia\e 
the  same  osiiiotie  pressure.  thi'V  are  sail!  to  lie  isiihiiiic;  when  that  of  one 
is  irreater  than  that  ol'  ihe  other,  it  is  A  v/ir  rhuiir:  and  when  less.  Iniimhiiilr. 


Biological  Methods  for  Measuring   Osmotic  Pressure 

.\  piaetieal  hioh.eieal  appli.'atioii  o|'  these  luineiples  rail  Very  readily 
!„■  made  if.  iiistrad  ol'  a  ri-id  semiperineahle  meiiil.raiic  sueli  as  Ihat 
li,_Mir,'d  in  the  dia-ram,  we  employ  one  that  is  esleiisiMe  and  takes  the 
t'oriii  ol'  a  i-losed  sae;  then  as  dilTusioii  ol'  water  oceiirs  llic  sac  will  either 
disteiiil  wIm'Ii  il  contains  a  stronevr  solution  than  that  outside,  or  shrivel 
,,!•  creiialc  when  the  ivvi'i-sc  .•oiiditioiis  oMaiii.  .Miiiiy  aniiiuil  and  veu' 
etahlc    protoplasmic    iiMMiiliranes    arc    sciiiipcriiieal.lc.    includins;    tlu'    <'ii- 

vclopc'  of   red   M I   .•orpiisclcs.     •riiiis.   if  w  .■  exaiiiine   blood   corpuscles 

iiii.hM-  the  micros,.ope  a'  I  add  \"  them  a  saline  solution  of  hi-rher  os- 
motic pressure  tliaii   hhmd  s.riim.  they   will   visibly  diminish  in  size  and 

l,cc irremilar  in   sliape;   \\lic>reas  if  the  solution   is  of  lower  osiiiotic 

prisMire,  they  will  distc^nd.     If  no  chance  ■icciirs,  the  osmotic  pressure  .d 

tl,,.  rell  I tents  must  c,|iial  that  of  the  saliii.'  solution  in  wlihdi  th dls 

an'  immersed,  from  which  it  is  clear  that  wc  can  readily  det.'rminc  the 
iiiauuitiidc  of  the  osmotic  pressure  if  w  c  l<iiow  the  streiiijlli  of  llic 
saline  solution. 

Instead  of  iiieasiiriiii:  the  iiidi\  idiial  cells  under  the  microscope,  wc  can 

asiirc  the  space  they  oci-ilpy  in  the  lluiil  in   which  they  arc  siispcndc.I. 

For  this  purpose  a   iioition   of  the  suspension   is   ph I   in   a   itraduated 

tube  of  narrow  bore,  which  is  rotated  in  a  liori/ontal  iiosition  by  a  ccii- 
trifiiire  after  bi^iiiv  chased  at  one  end.  The  eraduatioii  at  which  Ihe 
ui.pi'r  edce  of  the  column  of  cells  stands  after  cent  rifucinir  is  a  iiu'asurc 
of  the  relative  aiiioiiiil  of  cells  and  lluid  ill  tlic  suspension.  ITavini.' 
found  this  \aliie  for  <-e!ls  suspeiiilcd  in  an  isotonic  solution,  as  for  blood 
corimscles  in  blood  scriliii.  we  may  then  lu-ocecd  to  ascortain  it  for  the 
same  cells  suspended  in  an  unknown  solution;  if  wo  lin<l  that  tlio  cells 
oeciipx-  a  irreatcr  volume,  the  saline  solution  iiiiist  have  an  nsmotie  i)res- 


i,\\vs  or  soi.iTioN 


sMif  tliiit  is  lower  thiiii  lliiit  ci"  scnmi  in  iiiipnixiiiialf  piopniti.m  Id  llic 
rt'iidiiiL:     111!  Ilic  liilic  ill  tlic  tw"  i-;iscs.  anil   r'n    r<  rsii. 

Tlif  al>civ('  ai)|>!iriitiis.  ealliMl  a  liniiatoci-itc  I  l-'iir.  "J  I  lias  !)crii  v.-iy  t'\- 
tciisivcly  used  in  the  (M.ll.Tti..n  .if  .lata  .■.Mii-cinin-;  liic  rclativf  ..sini.ti.' 
pi'.sNiirrs  (if  tlilTfi-.'nt   i)liysiiilii!.'ii'   tiuiils. 

Hemolysis 

AniitliiT   wav    fm-   .l.'ti'i  iiiinintr    tli.'    niativ.'    dsnnitic    picssiin-    nf   .lif 
rrrcnl    siiluliiiiis    ccmsisls    in    placinii'   ('i|iial    aiinuii  ts    la    d-w    .Inips)    .if 
liliHi.l  in  a  sciics  .if  t.'sl  Hilics  .•..iitaiMiii'.'  Milutiims  .if  .lilTficnt  sti'«'n<_'llis, 
and  after  all.iwin-  \\u-  Inli.s  t.i  staml  fur  s.iiiie  tinn'.  mitinu:  in  wliieti  <if 
111. Ml    lakiiiji    .if    ill.'    Iil.i.id    e.ivpiisel.'s   ...•(•MIS.      Ill    s.iliiti.iiis    wliieli    are 

iMil.ini.'    (iv   liyperti.liie    "itli    tlie    cillteHts    ..f    tli ipilseles,    the    latter 

will  settle  t.i  tlie  li.ilt.ilii  .if  III."  tiilie  ali.l  til.'  siip.-riiatant  llilid  will  li.' 
Miiliiited  with  h.'iii.n.'l(iliin.  Imt  in  s.iluti.ms  which  are  dislin.^tly  hyp.it.iiii.', 
Ih.'  s.'diiiieiit   will  li.'  I.ss  .lislin.'l  and  th.'  Mip.'inataiit   llni.l  r.'d. 


Hy  n.itiii','  tl>  the  lowest  .•one.'iitiati.ui  i  per.'enta!.'e  .'o:  position  i  <d 
the  solutions  in  whi.-li  th.-  .•orpusel.'s  sink  t.i  tin'  liotl.n  and  h'ave  th.- 
supernatant  tliii.l  eolorl.'ss.  an. I  C-M  the  liiu'lf.'st  coneintiation  in  wlii.-h 
the  e.iriiusel.'s  when  they  settle  l.'av.'  the  sup.'rnatant  llui.l  r.'d,  we  i-aii 
deterniiiie  the  liiiiitiiiy:  eoiicentiations  f.ir  s.ilutions  .if  dilTerent  siih- 
stane.'s.  Thus,  with  liulloek's  Id.iod  th.'  followiin;'  r.'sults  w.'re  olilained 
(llaniliurjrer) : 


SlHSTANCK  PK.U.KNTA.iK.  Snii  M.ril    (IK  Si  M.rTKIS    IN    Will.  II: 

I  II 

.si-i'i;i:N.\r.\NT  Ki.rin  sm  iinm'.nt  n.rin 

WAS  Clll.lllll.l.SS  U  AS  l:'  11 

KNO,  I.IH  ii.'.i.'i 

Xii<'i  ii.Ciii  (i.'ii; 

K.SO.  I.Ki  t.liii 

<',  li.o,,  (riiiii'  siii;:ii)  (!.:.".(  ."i..;:; 

ClICOOIl    I  I'.it.   :i.ot;it<')  1.07  tm" 

.M«s(),.7ii:;i)  :!..'■:;  :■..:;() 

CaCI.  (l.s.-i  li.7!l 


3U'  ,v 


n  rin  sii  imiii:mii' \i.  nvsis  up  rinsiin  ncir  \i,  hhm  issi.s 

'I'lir   iiifjili  111'  llii'M'   liiiiithii;  ciiiicciil  I'jil  inll^  is  ilic  (■/■/  uiifi  ii  I  rut  mil 

iiiiil   ihiliciilcN   tlir  slrrh'^tli   lO'  fiirli    Miliitinii    llial    I'lili  lili'il    1(1   lildiiil 

williiilll  i-iJllsiliLT  iili.\  iliiliKi'^i-  111  1  lie  i-ii|-|iii>rl('S.  'I'lii^  <'lili'-;il  nuict'll 
Iriiticiii  is  iml,  as  iiiit;lil  al  liisl  siulil  lie  iiiia'j:iMi'(l,  tlir  same  as  llial 
wliirll  is  isdliiiiic  willi  lite  cnlitilits  uf  llir  (•(ii|iusclcs,  Lilt  ilistir,rll\ 
liclciw  il.  Till'  ii'asnii  I'lir  lliis  liri-iiiiics  a|i|iafiiil  il  w  i'  hIisimnc  iIic  lir 
lia\  iiU'    111'    ciiiliusrlrs    slis|ii'lii|ril     ill     all     isntiiiiic    siilllliiill     wiiicll     is     llini 

Lrrailinll\  ililiilril.  As  ililiitimi  |iiiici'iils.  ilir  rm  imscli's  liistcinl,  until  ;it 
last  lli<-ir  riiN ciiiiics  liiii'st  iiikI  llir  liriiiiiulnliiii  is  ilisfiiaiiii'il.  Tiic  liiii- 
itiiiy  (•iJiiiTiiti-aliniis  (iT  a  yi\i'ii  sail  \a!\  I'm-  ilirrciciil  i-iir|)iisi-li'-;;  llnis, 
tilt'   rnlicrlil  lalinii    ul'   siii|ii|iii   ell  |i  i  iji  Ir    siilulinii    lliat    just    railscs    lakill'^    lit' 

I'lii'^'s    iijiMiil    i-iii|ii|si'|(  s    is    II. "Jl     pel-    ri-lil.    111'    lillliiail    111 1    0.-17    |ii'r    iMIit, 

ami  111'  liiiisi'  lilniiil  u.i'.s  |i,.|'  i-i'iit.  It  is  till'  slifii'ilh  111'  till'  (■iii|iiisiMilar 
fii\clii|ii'  rather  than  vafiatimis  in  thr  usuinlir  in-rssuie  ul'  lln'  ciintrnt^ 
thai    is   ris|iiiiisililc    I'm-   llicsi'  ilirrrrciH-is. 

Till'  aliii\t'  ili'scriliril  iiirlhiiil  ul  lii'iiinl,\  sis.  as  it  is  raMnl,  I'liii  nol  In' 
ilsnl  r.ii-  i'iini|ia  risiiiis  ,,(  n^niuli.-  |irissiii  r  in  casrs  in  wliirli  llic  suliitiun 
ciintains  siilislanci's  whii'li  ailrr  tin'  lu'niii'aliilil y  ul'  llic  (•(ir|iusciilai' 
(■n\  i-lii|i ;  I'll!'  i\aiii|ili-.  il  i-an  iml  lir  Ilsnl  wlh'ii  iiica.  ur  aiiniiiiiiiiiiii 
sails,  111-  i-i'itaiii  liiNii-  liiMlics  ai-i'  prcsi'iit.  'I'liis  \i'<-\  I'act  is,  hnw  ex  cr, 
put  111  a  llscrul  piiipiisi'  in  ;isiTit;iinilii;'  ulirlhrr  a  uixi'li  sulistalirc  ilncs 
liavi' a  ilaliiaLiiii'..;'  iiilliicin-r  mi  tin'  ciirpusi-niar  rnxrlupi'  hy   tiinliii'j:  whi'tlirr 

hciiiiil.x  sis   iirs   when    we   siisprml    Ihr   curpiisclcs   in    a    snIiitinn    that    is 

isiitiinii'   with   thr  ciirpiisi-iilar  I'miti'iils.      We  i-aii    I'lirthcr  ili'tcnniiii'   tlii' 

<l('^n t'    this    |ii\ii'     inlliHiicr     liy    est  iinaliiiii     iiy    cnlin-     ciiiiiparisuns 

(('(iluriiiict  r-.\  the  aiiiiuint  nl'  hrniiiuliiiiin  that  lias  ilirfiisi'il  mit  uf  ihr 
porpilsclis. 

•     Plasmolysis 

An  analn;;iiiis  uii'tliml  I'ni-  ili'tirininiirj'  nsiiintii-  prrssiiic  is  that  nf 
plasiiinlysis.  in  whii-li  the  lirlnix  inr  ul'  certain  plant  eells  is  nliseiv  eij 
IIlicniscii|iieally  while  the\  ,ile  in  emitaet  willi  snluliuns  ul'  ililTerellt 
streipj-ths.  When  the  sm  iniinrlin'j  siilntiuii  is  isutnnie  willi  the  eell 
eiilitelits.  the  laltei-  lill  llie  erll  aliil  e\teiii|  lip  t.i  the  iiinre  ur  less  ri','ii| 
eell  wall  '.I  in  h'ii;.  :,  ;  luit  when  the  sululiuli  is  hyputunie,  the  ei'M 
eulileiits  lieeuiiie  detaeheil  lii  ill  the  cell  wall  at  uiie  ur  linn'e  places 
phisniulysis  ill  allij  ('  .  The  seiiiiperiiiealile  iiieiiililane  in  this  e.ise  is 
therel'iire    nut    the   I'lil    wall    Imt    ihe    laxer   ul'    p|iitupl;isiil    uii    the    surl';ice 

"if   tl II    eiinlelils.      The    iiiethuil    call    he    usei|    un|y     fur   del  ect  i  11 LT   sulii 

tiuiis  tliat  are  jupertunie.  !••!•  >>.  ilh  thuse  th.-it  .-iie  !i\  pntmiic  tin-  ee!K 
merely  lieeunie  iiir'.:ii|  ami  e\irt  mure  pressure  un  the  mure  ur  less 
ri'jiil    cell    wall.      .Maii\    ul'    the   einielusiuns    thjit    lia\('    licen    <lr;i\\ii    rruiii 


I.\\\^    ul'   r-(PI,l    rlll\ 


•) 


I'i'Mills  iilihiiiM'!  Iiy  llii'  |il;iMii<>l\  I  ic  iiirlliixl  li:i\<'  ico'iiilv    I n  cnlli'il   in 

(jiiol iijii,  lii'c.iiisc  Mil  n'ir;ii(l  li;is  lni'ii  t;ikcii  ul'  llir  |iii\\cr  nl'  ihc  ciilluiils 

.i|'    ihr    l-l'll     til    illlMM'll     '    illlllilll'  I     U:ill|-     1  M  r    IiM'JT    (i'J '. 

Till'  lllctliiiils  lit'  lnii]iil\  sis  iiiiil  pliisiii(il\  sis  li;i\('  licfii  llscil  liii-  till' 
invcstjij.-il  inn  111'  rininv  pinlilciiis  in  ninliiMnc  In  llir  c.-isi'  ul'  I'ciliiin 
tnxir  lluiils,  surii  ;is  snjiLr  xcnn.ii,  Iclanns  Insin.  cliv.  ilcli'i'ininjilinn  ul' 
llir  liriiiiil,\  til-  |iii\\ri'  lias  |iiii\ril  iij'  \iiliic  in   iiiiiiiliiy   assiivini;  tin'  dam 

a'^inir  inliin'M n  nllii'i'  cfils  tiian   lilninl  i-iir|)nsi'lfs.     Stmliis  in   licitiol- 

\  sis  liavi"  alsii  liiTii  i's|)''i-ially  \aliialili'  in  wnfUini;  mit  tin'  irncliiinistn 
liv  uliirli  I'i'lliilai'  toxins  in  i.>'<'i>''>'al  ili'\i'l<i;i  tlirii'  arliim.  ami  tlii'  rnmlitiniis 
nnilri-  wliii'ii  lliis  acliiin  may  In'  cimntiiai-li'il,  as  \>y  llir  il('\  I'lnpini'iit   of 


'f  ■■■  X 


:r 


.( 

s 


li  ' 


A 


it 


\'~-     .!.       !■"      -li.iu      i.;,,-in.,lv-i-      ill      .,i;.      11. .in      Tr.i.l, 

l-.i-"'"K-i-     111     II    ■  '     M       -iiLv.;  .     I'.     I ;kui.I     I  I.,- 

"■il'l   /'.    llii     |ii.il"|'l.;-ii,,       I  An.  1     ill     \  n.  -  I 


,li-,..|,.i.         I       iH,,iii.il     ..11;      /; 
Ul      l.,l     .\1.      kNn,!;     h.     llu      I.  II 


anliliii<lics.  I'uitlirniiiiic,  an.\'  snliitimi  tliat  is  to  lie  iiijcctcd  info  tin' 
animal  lii>i|>,  citlii'i-  iiit  ia\  (■ninisly  ur  siilnMi1ani'iiiisiy,  sliuiilil  (ifst  of  all 
lie  Irstcil  l).\  the  aliiivr  mrlliuils  in  iinlci-  tn  liinl  mit  \\  lictlicf  it  is  isnluiiii' 
witli  till-  liDily  Ihiiiis.  If  a  li.\  pi'itonic  siiliitii)ii  is  inji'i-tcii,  it  will  result 
ill  llic  alisti-actinii  of  water  fi-nm  tlic  tissito  ci'lls,  wlicicas  a  li>  piitnnic 
siiliition  will  caiisi'  tin'  watiT  cunli'iit  nl'  tlii'sc  to  inrrcasf.  Advantairt' 
has  I'cci'iilly  ln'cn  laUcn  nl'  tins  w  alcranstiai-tiit!,'  rtfcct  oT  liy|)i'rt<)nic 
snhitions  in  tin-  lical  nii'iit  of  \M)iini|s.  |!y  i-onstantly  liatliinir  tlicni  with 
striiii'j:  s;iliiu'  snliit  imis,  an  niillluw  nf  wal<'r  is  set  up  frnin  the  tissue 
<M'lls  thai  liiuili'i-  nil  the  wiiiiinl.  anil  this  teinls  In  hrinir  tn  the  rnclis  (if 
inliM'tioii  the  ili-t'ensi\e  sulislaiiees  that  are  presi-iit   in  aninuil   tluiils. 


illAl'TKK   II 

OSMOTIC  i'i.'i:ssri;!';  iCintMi 

Measurement  by  Depression  of  Freezing  Point 

Tlu'  liiiiilMtidiis  in  tiu'  use  cii'  the  iiliiMiinlytic  aiiil  licin<il.\  tic  nictiiods 
ill  tlic  pii'cisc  iiH'iiMiii'iiu'iit  i.f  till'  osMiiitic  pii'ssiiri'  of  till-  liiidy  llniils 
li.iNc  ri'iiiliTcij  it  iiii'cssMiy  tn  liml  sn;ii('  jiliysical  iii('tlnni  tli;it  will  lie 
aciii'i-iilly  iiplilii'.-ilili".  liccMiisi'  (if  tccliiiiriil  (linii'iill ii-s,  it  is  iiii|)riirtii'iilil<' 
til  iiir.'i  .iiif  till'  pii'vMUi'  ilirirtly  liy  I'liiplnyiiiLr  :iii  iisiiiniiii'trr.  so  tliiit 
sciiiii'  iiiiliii'i't  iiii'tliiiil,  (li'pi'iiiliiijr  nil  ;i  iciiijiiy  iiii'iisiirjililc  jiliysiciil  ])ni|)- 
iity  wliicli  \iiiirs  ill  piiipiiitiiiii  to  tlic  nsiiiiitii'  ]>i'ossuro  iif  the  (iissolvcil 
Milistiiiii'i's,  iiiiist  111'  iiscil.  I'"(irtiiiijit('ly,  one  siii-h  exists  in  tlic  jiniperty 
wliicli  ilissii|\  111  siilistiiiii'cs  liinc  ill  low  criiiLT  tlio  tt'iii])('r;itiiro  iit  wliicli 
I  III'  pure  MiKciil  Miliililic's:  till'  fici'/iiii;  point  of  ]iur('  wiitcf.  for  ('x;iiii|)l(\ 
is  iowi'ii'd  wlnn  siiiistiiiicis  arc  ilissolvcil  in  it,  and  the  extent  of  this 
lou  I'liiiir,  with  ccitniii  ri'sciViitions  which  will  li-  explained  later'  (pane 
111),  is  propHiiiniiMl  to  liic  iiiolcciilar  concent  rat  ioii  of  the  solution  and 
iiiiicpcndciil  of  the  cliciiiical  nature  of  the  sniistance  dissolved.  This 
lo\M'iiii!.'  of  tciiipcialiiie  is  dcsiirnated  liy  the  (Ireek  letter  A,  and  to 
miasiiie  it  a  lhci-iMoiiictei'  is  nsed  which  is  not  only  extrciiidy  sensitive 
liilt  in  vhicli  the  level  of  the  mercury  colnniii  can  ho  adjusted  so  that  it 
stands  at  a  coinciiicnt  level  on  the  scale  correspondiiifl  to  the  freozinn 
jioiiit  of  \\liati\cr  sii]\eiit  was  used  in  making  the  solution  under  investi- 
L'ation  I  Ucckinaiin  "s  tlicrnioiiictcri  (Fiir.  4).  The  exact  position  on 
the  scale  of  tills  tlieriiionictcr  at  which  the  ]Miro  solvent  freezes  havinir 
liecn  ascertaiiieil,  the  ohscrvatioii  is  re|)eated  with  the  solution  whose 
osinotic    ;)iessiire   is   to   he   detei'iiiined. 

A  uiaiii-iiiolecular  solution  in  water  (Inn'iiir  thcroforo  nil  osipotie  pros- 

sui f  17(1. 240  mill.  H'.'i  has  a  frceziii<r  point  that  is  l.StP  ( '.  lower  than 

that  of  |)iirc  watci.  This  is  known  as  the  "freezing  imint  eonstniit." 
and  it  varies  for  dilTcrcnt  solvents,  liointr  ■!.!•  for  acetic  acid  mid  4.0 
for  licii/.ciie.  If  an  unknown  watery  solution  is  fouiui  to  liave  a  frocz- 
inp  point   that   is  A     f.  low.r  than   that    of   water,   its  osiiiofic  pressure 

u  ill  eipial        ,   , ,        mill.  Mi,'. 
I..^t) 

10 


OSMOTIC  ri{i;ssrKi: 


11 


Till'  ili'|)n'>siiiii   III'  I  III-   i'l'cr/iiii;  points  pripdiicccl   liy   tlic   viiiidiis   licidy 
lliiiils  liiis  l)ccii   <'oiii|iiiii'(l,   tlic   objects   in    view   lipiiit;   to  see   whether 

oslllcitic  pi'csslirt'  is  ii  property  ullieli  eluiimes  limlei'  dilVefeiit  pli.\  sioloirie 
.Hill  p,illio|(p<rie  eoiKJiiions,  jiinl  to  find  out  1)\  eoinpiiiisoii  of  tlie  osmotie 
pressures  ot'  the  lluids  in  eont;iet  with  m  iiieiid>riiiie.  wiiellier  jiliysiejil 
t'lirees  ;iiolie  ciin  lie  iiehi  I'esponsilile  for  the  t  riinst'ereriee  of  snlistnnees 
tlii'oiiirli  it   I'roni  one  llnid  to  tl tluT. 

The  Rolv  of  Osmosis,  Diffusion,  and  Allied  Processes  in  Physiologic 

Mechanisms 

An    iieeount    of   some   of   the    in\  esliiriit ions    in    wliieii    the    I'ori'U'oint; 
nietiiods    liave    iieen    used    will    iilustnite    their    viilue    in    revealinj'    tlii' 


^' 


\ 


n 


r 


1■■|^■-  4.  Api  ;.t.ilus  f.ir  nu  a^unriuiit  c.l  the  (U-|>ri>si..ii  of  I'ui/iiiK  i^'iiit  I'l  ^'.liili.ii^.  The 
'.ilun..n  IS  |il,i..il  hi  tho  laiKL'  u^l  lulu.-  vuUi  tin-  m.Ii-  artn,  :ii>>l  iii  il  m  mi-|'<ui|i.|  I'a-  l.ulli 
■  I  .1  llitkuuinn  tliiriii'iiiu-tcr  with  a  iilrniiiiini  l.mp  to  >,  r\f  i..r  stiiiiuK.  'I'lii-  virpi  i  cnii  ..t 
tlu-  iiuuor>  inliimii  ,,t  the  tluTiiionKtfr  i>  ,s]|m\ui  lii:i.;iotM-.l  at  tin-  m-rer  Iftt  eoiiiir.  'lllo 
am. .not  ol  iiurmiv  in  the  tluriii.mu-ttT  tulir  lai;  In  nuiiiatf.l  by  tappiuK  ll'e  tiri't  r  tin!  null 
tti.  tin  niiMiiKUr  111  various  povitinns,  Tlie  tt-l  liil.r  is  piotcitt.l  liy  an  uut.r  lultc,  \\tiKli  is 
llivn    i.iatr'i    ill    a    vis&tl    containiny    a    frccziiiK    iiustiiri'. 


nieelianisni  involved  in  the  transfeu'iu-o  of  water  and  dissolved  suh- 
stanees  through  eell  memhranes,  as  oeeurs  in  absorption  of  food  ill  the 
intestine,  in  the  fonnatiim  of  lyiiipli  and  urine,  and  so  fortli.  In  cni- 
ployinsj  ])hysieal  niethtnls  in  tiie  .aieidation  of  siieh  problems,  it  Is 
always  most  necessary  to  proceed  with  great  care,  since  the   i)hysical 


u 


I  ll\ -II  iM  III  Mil   \i     I;  \>l.'-  Ill'   im  >iiil  iii.KM,   luiMissi;: 


i'liriin>I  vviiiks  willi  |iilii-  ^ciliil  11.11^,  wlilli'  ill!'  |ili\  Niiilii'_'i>l  liiis  111  il>i' 
lIlliiK  IIkiI  ;ilr  jl«:i\s  riini  |il  ii-,i  I  ril  Mini  riri|lli  III  I  \  \i\\  \  aiijllilr  ill  i-mn 
|iiisil  lull.  W'l'  iiiii^l  Niih|ilii\  till'  |iiiili|riii  ii>  i'jir  ;i>.  |)iissilili'  liy  li;i\  iii'/ 
I'liilll.v  111  lull'  lis  llii'  rsMi-1  IMllllr  uT  llir  I lii ili i'-'ir;i I  |ili'lili'lll  wllii'll  .'I  fiilll 
liMllMili  111  |'li,\  ^il■ll^ll^llli^.•ll  \.ililis,  Midi  UN  usliliilii-  |il(':  lie.  Ml,  y  rllll 
lilf  lis  111  rhii-nhili'.  iiiiil  w  1'  iiiiisl  I'liiisiilri-  llic  iillicr  |iliysic;il  t'oiccs 
wliii-li  iiia\   .■is'-is|  III'  I |ir\   ihr  |i,irl  iciiliir  mil'  \\i-;irc'  iii\  csl  iciitiii-^. 

Ill    llli'    |ili\Ni.M|    r\  |iil  illlcIlN   ill  >i-liliiil    illiiiM'.    Illi'    srll|i|)iMllll'.lli|l'    llirlll 
lilMlir    lii:i,\     ill'    rii|irri\  I'll    nf    ;is    I'nlii  |iiisi'i|     iiT    |iiiii's    nl'    sUrll    M    si/i'    lli.ll 
lliry     |iiiiiiil     iiiil.\     llir    siii.illrsi     iij'    iiiiilrrillfs      llm-i'    iij'    wjilt'l'      In    |i;ins 
lliriiii'jii   llii'iii.     Si'iiii|iiiiiir;ililr  iiii'iiilii'iiiii's  \\illi   l;iri;ri'  purrs  iiiav .   Iiuw 

I'M'I',    csisl        lllill     is.    IIKiMlll'.llH  s    (vllirll    |illinil     W.lIlT    llluill'l  I  It's   illlll    lliull' 

rilli's   of   siiiiplc   cliniiii'al    siijislaiici's   lu    pass,   Iml    Imlil    liark    llms in 

puscil  iif  laii;r  rmiiplrx  iiiuiri-iilis  Siicli  a  srniipiTiiiralili'  liicmliialii' 
VMiiil,!  alluxv  ihr  saiiiii-  I'uiisl ii  in'iils  Iml  imi  tin'  pruii'iiis  uj'  liluuil  sri'iiiii 
III  pass       ll    is,  liu\\c\i  r.   III!  Iuiiv;i'i'  siniipi'niii'aiili'  luwai'ils  all   uf  tlir  ilis 

suUi-il  siilislaiii-cs,  aihl   llii'   pr -ss  ul'  (lil]iisiiiii  I  hiuii'jli   it    is   iiiurc  '^ciu'I' 

all.v   ilrsiM  iialiil  as  uiic  ul'  ilmhtsis  lliaii  ul'  usiiiusis. 

SiiiiT  ilir  passa'jr  ul'  ilissiih  I'll  inuliiMilfs  l|iruii!,'li  iiu'iiiluaiii's  ih' 
pi'iiils  iipiiii  llii'  piiiiciplc  ul'  iliri'ilsiuii,  lis  lali'  will  lir  piupurt  iuiial  In 
till'  usiiiulii'  pii'ssiiii's  ul'  till'  siiluliiiiis  nil  till-  two  siirTaci's  ul'  ihc  iiii'iii 
'iiaiii'  ami  In  Ilii'  •>]/.<■  ul'  tliv  iiiuli'i-iili's,  small  muli'i-iili's  ililTiisiinr  inui'i' 
ipiirkly  lliaii  lari;r  inns.  Siippusi'  a  iiii'iiilnaiic  pi'i'iiii'alilc  In  siiiliuiii 
clilmiili'  ami  walm'  is  plai-iil  lulw  nii  two  lliiiils  I'uiilaiiiiiiii'  suilinm 
I'lilui'iilr  ill  siiliiliiiii.  lull  in  ^ii'i'ati  r  i-um-i'iit  raliuii  in  mii-  ul'  llii'in  lliaii 
ill  III!'  ulliiT:  tlir  suilimii  I'lilui'iili'  will  dilVusc  I'luin  llic  st  luiiu'ci-  In  llic 
wi'aki'i'  suliitiuii,  am!  wain'  will  ililTiisr  still  iiiuii'  ipiirkly  i  lii'i-aiisi-  its 
iiiiili'iMilrs  arr  siiiallci''  in  tin'  uppusiti'  iliri'ct iun,  until  tin,'  iiniiiln'i'  ul 
sniliiiiii  i-liliu'iili'  iiiiili'i'ulrs  in  a  uIm'ii  \uluiin'  uf  suhiliuii  is  npial  un 
liuili  siili's  111'  llic  incMilii'am'.  I''iii'  a  linn',  ihri'ct'un'.  tlii'  vuliiiiii'  of  tlir 
st  i'un'.:i'i'  sulutiun  \\  ill  inrti'asc,  Tlii'  ililTcii'iicrs  wliirli  rsist  in  tlic  ilif 
I'lisiliilitv  nl'  (lissiih  I'll  muliTiilcs  ai'i'  aiialnLTuns  In  tliusi'  wliirli  liavr 
Iniiu'  lii'i'ii  kiiiiwii  tn  i'\ist  ill  III"  ililTiisiliility  nl'  jiasi's.  luit  till'  I'l'lalimi 
hitwi'iii  lair  ul'  ililTusjIiiliiy  ami  iiiuli'riilar  \M'ij;lit  is  nut  su  siiiipli-  as 
tlir  raliu  lii'twi't'ii  tlii'sr  t'\u  ipianlitirs  in  i;asi's.  Tlii'sc  ri'latiinislii|'s. 
ImwcMT,  iinlii-atc  scviial  iiutliri-  pnssilulitii's  in  tlic  rxplanatiun  uf  tlir 
iiii'i'liaiiisiii  uf  f\i-liam.'i'  uf  sulistaiii'i's  tliiiuiiili  iii('iiilu'aiii'><,  anil  must  iiul 
III'  u\  I'llnnki'd,  as  lliry  uftiii  an',  in  tlir  inti'ipri'latiiui  uf  i)li,vsiulnirii' 
lilicniiiiii'na.  An  r\fi'llrii1  rr\ir\\  uf  tin-  pnssilili'  I'liiiiiitioiis  is  jrivi'H 
li,\  Stariiiii.'  ill  ilis  "limiiaii  l'li,\  sin|n!.r\ . '"■  j-'nf  cxaiiipii',  let  us  suppnsr 
till'  snlislanri's  mi  the  twii  siilcs  nf  a  si'iiiipcinii'iilili'  iiicnihraiic,  siicli 
as     till'     pii'ltunral.     In     In'    ililTiTiiit     in     ililTiisiliility,     as     raiic    siiirar. 


osMdTir   i'His>ii(r;  1'! 

uliic'li  <ln.'s  lint  i.'ji.lilv  .lilTiiM',  Jin.l  siidiimi  fliloii-lc,  uliirh  .lilTus.'s 
.Hiickl\  :  till-  nMiiotic  (low  will  Inkf  plii.-c  fioiii  llic  su.liimi dilorid  solii 
tic.n  t"  tin-  ciiiir  Mi'-'iir  rviii  wlicii  llic  s(i«liiiini'liloii.lf  Mvlutinii  is  siniiiiriM- 
tli.iii  tlic  Misriii'.  In  siii-li  a  i-iiM'.  wiit.T  iiiuli'iMilcs  will  p.iss  liimi  tin-  tluid 
lijiviiiir  till'  liiali'T  oMii.itic  ]Hrs>uic  I  NiiCi '  tc.wiinl  !i  iliii.l  ill  wliii-li 
iliis  is  liiw  IT  I  siijriir  . 

Fiirtlii'iiiHirc,  tlir  sinipii'  laws  i.i'  osmosis  iii.'iy  ho  ii|is.'t  l).v  mi  iiltrin- 
ti\i'  iiilliii'iKT  of  tlic  iiM'iiilniiiic  toward  certain  siilisfaiicfs  IdiU'  to  tlo'ir 
iMToinini.'  dissolv.'.l  or  adsotlifd  in  it  i  scr  pai;.'  ti.")  I  Imt  not  towanl 
oili.'is.  .Many  ni-'iiiliraiK's  of  tliis  nature  arc  known  to  ttic  eliemist 
(■.<;,.  rulilier  iiieniliranes  in  eoiitaet  willi  u'ascs.  pyridine  solutions,  etc.i, 
an. I  it  is  prol.aMe  tlial  siieli  a  pioperty  of  sideetive  soliiliilily  may  |ilay 
a  not  iiniiiiliortant  role  in  the  traiisfeii'iicc  of  siilistaiiees  acniss  animal 
iileiMlirani's    i  Kaldenliei'i;  ' '. 

Tliese  few  eonditions  wliiidi  may  modil'y  tlie  direction  of  tlio  osmotic 
llow,  all'  indicated  lure  to  sliow  liow  involved  siicli  problems  are.  and 
liou  careful  we  must  Ijc  not  lo  assiiiiic  tliat.  liecause  a  siilistaiice  is  traiis- 
feiied  tliroiiiili  ii  livinir  memliranc  contrary  to  tlio  simjilcr  laws  of  os- 
mosia  and  diffusion,  it  must  involve  tlie  expeiiditiirc  of  forces  dilTcrent 
from   tliose  operatiii','   in   dead  mcinliraiics, 

.\notlier  fore mes  into  opi'iatioii  under  certain  conditions     namely. 

Iliat  of  tillriiliiiii.  Tliis  is  a  piiitlv  mecliaiiieal  process,  in  wliicli  iiiole- 
eiiles  are  forced  tliroiiuli  tlie  iiores  of  a  liltcr  i  i.  c.  memliranc'i  liy  dil'- 
fcrences   in    pressure   on    its  two   sides. 

\Ve  are  now  in  a  position  to  consider  in  liow  far  the  almvo  jiliysical 
forces  exiilain  certain  iihysioloiiic  prolileiiis. 

1.  Is  III'  iiUsiifjtliiiii.  'mill  lilt  hliiiitl  mill  hiinfih  liri'iililliiiil  In  tlir  iiilf.<- 
tiiiiil  inills,  iif  siihuliniii  >t  in  Kiliiliini  hi  tin  lull  si  iliiil  rniili  ills  iiilinlif 
ill  fii  lull  I'  jiiiii  Ihi  iininssts  iif  lillniliiiii.  tllffiisinii  nml  iisninsis.'  The 
alisorption  of  weak  solutions  of  liii;lily  dilTiisilile  siihstances  is  iirolialily 
very  larirely  a  liialtcr  of  osmosis  ,ind  ditVusioii,  and  water  passes  ipiiidvlx' 
into  the  Mood  liecaiisc  of  osmotic  attraction.  Imt  that  otlier  forces  ordi 
narily  come  into  plax  is  \er>  clearly  estaldished  liy  tlic  followiiij;  oli- 
scr\  ations.  If  a  piece  of  iulestine  is  isolated  from  the  rest  liy  placiiii: 
two  liiratnres  on  it.  and  the  isolated  loo])  tilled  either  w  ith  a  solution  con- 
taiiiiii<r  the  same  saline  eonstiliieiits  in  similar  |iroportioiis  as  in  lilood 
seriiiii,  or  lietter  still,  with  some  of  tlie  same  animal 's  blood  serum,  it 
will  be  found  after  some  time  that  all  of  the  solution  becomes  absorbed 
into  tlie  blood:  the  contents  of  the  loop  are  therefore  absorbed  into  the 
blood,  exen  tlioinrh  llie  osmotic  |)ressnres  of  the  dissoUcd  sutistaiices  are 
tl  n  same  on  both  sides  of  the  membrane  i  Weymouth   Hcid''*. 

The   intestinal    menibranc  seems   to  |)ossess  towards    ivadily   dilTilsible 


II 


I'1I>>||  0(  IIIMK  Al,    HAMS   OK    I'llYMOIIMil 


<M.    l'Uiiri:sSI,.s 


siil.Ntiiiir..s  ii  |.,Mniiiil.iiil\  uhi.li  \iiii.>.  not  Ml  :ill  with  ilic  |.li>  sit-al 
.lilYllsiliility  ,,|-  i|„.  siil.Ntjiliic.  hut  with  its  Viilllc  Croin  a  phv  siolotfir 
Nlaii.||..iiiii  •|'liu>,  sMiliiiiii  sul|ilialc  ami  Mxliimi  .•hli.ri.lf  .lilTiisr  tlii<.u«li 
"iiliiiaiy  iii.iiil.iaiirs  uilli  aiiMiit  f,,iial  lai-ililx.  ami  \  .■!   if  a  sniiitioii  coil- 

taiiiiim   lli,>,-  t«..  salts   is   plarc.j    in   tli..   inlcslinr.   il hlniiilc   uilj   l.c 

alisnil...,|  iiit,,  til,.  |,|,„m|  „iii,.1i  mh.iv  i|uickly  than  tli.-  snlpliatf.  Sodium 
sul|.lial.  Hi  uatii.v  solution  .lilViis.s  llirou'jii  a  mnnlnanc  lilt,M-ii  tiiiifs 
rii"i"  'iMirlJv  than  raio'  suuMf,  l.iit  iiom  tlir  int.'stina!  lumm,  i-anr 
sMuar  Is  al.s,,ili.'.|  ten  tiinrs  lor.'  .|iii>'kl\  than  so.lium  sulphali'.  It', 
liou. 'Ncr.  Ill,,  vitality  of  thr  rpitli.'liiMii  is  .|.>sti-o\  cil,  as  l,y  liist  of  all 
I'athinu  it  with  a  solution  of  so,lium  iluoiiil.'.  thm  liir  sulphate  an<l 
I'hloiiili'   will   1p,.  alisoilM'.l  at   an  cijual   latf 

AlthoMvli  clil'l'iision  ami  osmosis  can  hot   tlicrct'ori'  play  any    si(rnilicanl 

i"l"    in    thr   mnriial    pio.Mss   of  al.soiption    fr the    iulrstiii.-.    \\v    must 

not  .nlif.'ly  ilisrount  th.'m;  i,nc|,.i-  .-.■rlain  <-in-uiastarM'fs,  these  pliysi.-al 
tori'.s  ma,\  ass.i  I  their'  inlluem-e  as,  f.,r  e\aiiipl.'.  when  eoiu'elltfated 
saline  solutions  aie  present.     Sm-li  solutions  will  attiaet  water  from  the 

''' '•   •""'•   '"'"'I'   Iliiii'-'s   lieins;  e.pial,    nn>re    will    I.e   attraet.-.j    the   less 

perm.aM.'  the  ipitlielium  hapjiens  to  he  towards  the  saline  employed. 
Sulphates    ami    phosphates    will    attract    iin.re    water    than    chlorides    or 

acct-tcs.     This   property   of  the  salim-  solutions  to  attract    water  i n 

terai.-;  !h.'  natural  tendency  for  llu'  water  to  he  ahsorhed.  and  the 
Iniue  vohuiic  of  Ituid  stimulates  peristalsis. 

-  /''/  //"  /,//</.s,V,//  i.roi;  ss,  .^  „i  jilt, ■III  ion,  iliffiisliiii  mid  osnio.si.i  siil- 
prf  I,,  ,ii,;nnit  fur  III,  i,n,il iirl inn  ,,f  iir'iii  h<i  llii  hliliii  i/s.'  I'mler  normal 
conditions  the  nioh'i-ular  cunc(>ntration  „(  :h,'  urine,  as  determiiu'd  hy 
the  depression  of  free/inji  point,  is  <-oiisiderahly  irreatcr  than  that  of 
''"•  l'l'""l  This  indicates  that  exerelion  iriust  have  occurred  contrary 
to  thi'  l,n\s  of  osmosis;  in  other  words,  that  the  renal  cells  nnist  ha\e 
'•"mpclled  dissolved  molecules  to  he  transferred  from  the  hlood  to  tlie 
nrinc.  althoii-h  the  diir.'renee  in  osiimlic  pressure  would  cuise  them  to 

1''''^-   '"    'I pp'isiic  direction.     This   fore.-,   sonu'limcs  called    f.u'   want 

of  a  hettir  name  --vilal  .ict  i\  il,\  ,•'  must  depend  on  the  operation  of 
in'..c,ss,->  that    are  .piit.-   distin.-i    from   those  of  .liffusion,  etc.;    hut    that 

""■>    ■■"■"  iieccsv.irily  of  a  ph.xsical  nature  i  c.  ir..  vital  i  is  less  prohahle 

than  that  thcv  dc^peml  on  sonn'  phxsical  process  the  natnre  cd'  which  our 
present   knowlediie  iloes  n^it   permit   us  lo  luiderst.-ind. 

liy    coniparinu'    the   osmoti,.    pressures    ,.f    urine   and    hlood,    attempts 
''■•i^''  '"■'■"  iii'i'l''  '"  measnie  the  work  done  hv  the  kidney  in  the  pro.lue 
'ion  of  .Hi!!...     Thus,  i-  has  heen  !„„nd  that  J>  :-.,i-  noruiai  urine  i  human 

is  ahout    I.N,  and    for  hi 1   .-ihuut   (l.li,   fr which   it   may   he  caleulate.l 

that   in    the   pi(,din-li(ui   of    I    kiloi;ram   o|    nriiu'    I.Vl   kiloorammetcrs   of 


osMiirn    I'lti.ssi  111 


15 


uiirk  iiic  t'XiH'iiilcil.*  liiit  lliiit  mhIi  fiiiripiii  isoiis  hI"  tlir  (iMiiiilic  prcs- 
•-iiii'  III'  IiIiiikI  ami  (iriiic  iiii'  t'iill!iciiiii>i  .'in  :iii  iiiiliriiliiiii  nl'  the  wmk  "I" 
liii'  kidney  is  ex  idcnci'il,  nut  iil'iiii-  liv  tlic  irMills  of  the  iilicur  I'liiciila 
liiiiis,  liiit  iilMi  li\  tlic  I'.'ii't  llial  iiinli'i'  <Tit;iiii  (•iiciiiiislaliccs  i  as  after 
I'cipioiis  iliuri'Hisi  till'  iiMriiiiic  jiroMiic  uf  tin-  mini'  may  lie  cniisiiliTalily 
Imrir  tiiaii  that  <if  llir  IiIikmI.  Tiiat  oppiPNiii'  ,  ,'laticiiiNliips  nIkhiIiI  exist 
iiiilieates  tliat  (lilTereiiecs  in  osindtie  pressiiic  iietueen  lilunil  anil  urine 
can   si^'ilify   little   if  anylliini!   rfualdini.'  tlie   work   done    li\'   the    kiilney. 

I'or  some  time  afti'r  tlie  applieation  of  osimilie  pii'ssurr  measiirements 
to  llie  Ntuil>    of  liiolou'ieal   prolilrms,   it   was  tliouuht   lliat    ileleiinination 

ipf  A  in  urine  miirlit  I f  elinieai  \alne  as  a  criterion  of  renal  ellieieney, 

esp.M'ially  in  one  kidiu'V  as  eompired  with  the  other,  i'oi'  this  purpose 
A  \vas  ileteriiiined  in  samples  of  nriiie  remoxcd  from  carli  meter  li\ 
eatheteiizalioii.  The  tests  of  renal  elVieieney  liased  on  the  rate  of  excre- 
tion of  |)otassiii)n  imlide,  phenolplilhalein.  etc.,  ha\c  however  lieeii  found 
of  much  greater  value. 

'■'<.  Is  llii  fiiniiiiliiiii  iif  hiiD/ili  jinnlii  II  iiliiislciil  jinniss.'  The  osmotic 
pressure    of    normal    lymph    is    neailx     always    somewhat    heiow     that    id' 

lilood    serum,    altliou'.di    oecasioiiall.\     it    has    I n    found    to    he    a    trille 

liitilicr.  rii.\sical  processes,  such  as  nitration,  nnnlit  tlierefoie  sllHiee 
to  account  for  its  formation  under  most  conditions.  Hut  when  w c  con- 
siijei'  the  esecssi\e  |iroiluction  of  lymph  that  occurs  as  ;i  result  of  cel- 
lular activity  or  followinjr  the  injection  of  certain  sulistaiices,  called 
"  lympliairoi-'ucs."   it    is  not    so  easy    to  explain   the   production    in   stich 

teriiis,  altholl'.'h   some   interest  in<,;  .'ittempts  ha\e   I n    made  to  do   so   liy 

those  that  jirc  wedded  to  the  mechanistic  view.  I'lir  example,  the  vei'y 
mai'Ued  iuercase  in  lymph  Mow  wiiiidi  occui's  as  a  result  of  muscular 
exci'cise  or  y;landular  activity  has  In  en  attrihuteil  to  the  fact  that  dur- 
ini;  such  processes  lai'trc  molecidcs  Ipccoine  nrokeu  down  into  small  ones 
in  llic  ceil  protoplasm,  so  that  the  osmotic  tiressurc  is  laised  and  water 
is  altraeted  into  the  the  cell  until  the  latter  lieeomes  distended  and  a 
jtrocess  of  (ilti'alion  into  the  neitrhlioiin!;  lymph  s))aecs  oecuis  (see 
pa^re  I  if)). 

There  are  sr\er,il  other  physioloudc  pioees.ses  of  secretion  ami  excre- 
tion which  miulit  l>e  considered  in  the  present  relationship,  hut  the  aliovo 
instiinces  will  suflice  to  illustr.ite  the  tfener;d  i>rinciplc  upon  which  all  id' 
them  have  to  he  considered. 


/ 


11-111. ,11,-   j,r.  ..viirr   .  ..i  r.  .i-nnlii  i;    1,,     ',  (M,-    l".   ,-,|„aK    ^.U'J   mm     tin    <r^    in     ..f    It   I  >l     ;iii.| 

•'•■''  '"rr.-s;...:!.!,..,;   I..    A  :   ■■  ■    r     .,;..,!.    i<,j.^..    „„„.    H^    ,JJ=   ,;,.    il.r,.       Tii,     .UlU >     .,    ll.,  ,. 

I"rr   ciiiial   ti,  .-1  cliimii   ..t    «:il,r    15(1  in.    hi«h.      Ai.or.lim:  to  thf«.-   .■al,iil.ili,,ii,    il    vi..ul,l   :,].y,.n   lli.it 
lli<-   ki.liuy   II)   |.i,i,!ii,ini.   till'   .-ivirii;!-   .l.iily   <iiit|.iit   ..I    l^mi  ,-,i-.    nriii.-  i.,rl..riM«   J.'i   kil"k'r.iniin.  t- r-^   ..f 
«urk   III   i..iii],.-in-.iii    with   tin-    U.iinil   kil,.KraMiMi<lir>   wliiih   tlu'   lu^irl   is  luii 
>.iini-    tiiiir    liiaiii-   Jl.'J. 


Mini>uti''l  t<i  iKriorni   in   lli 


il 


(  ll.vi'i  i:i;  III 
i;i,i:t||;|i    I  iiNJin  Tl\  ri'V,  IH^sim  l  AIIuN.    \\|»  |u\|/.\l|it\ 

■rilc     i.>hM.lir      |Mi'»M|l<'     U     nnl      i  II  II  !■(  |  l|r||  1  I  \       tnUllcj     \i>     I"'     .Mmsidnil  I 'I  v 
..'li'.'ll.T    lli;ill    lli;it    .  \|Mrlr,|     llolll    llli'    sIl.MMtll    <<i    tllc    s..llltic.ll.        Alll Ull 

ji  ,,f  ;i  .JIM  II .Icriihir  \MiIiT\  s.ihili r  .-a  111-  sii'.-ar  f  ''\1  um.  I"  lli''  li'''"'' 

is    I  M;     iv,-,.    |,,t.jr     l<>    .    Ih.ll    '<\    '...lllllll    clllniLli'     '■>>.'>    VHI.    U<    the    lilrr)     is 
,-nlls|,|.l:ililv      -iral.r         IT    IIm'     Ii.v  |...t  lirMs     l  rua  I  .lilP.'     llu'     Tflal  iMM-.lli|.    I" 

iiM.li.-iilar  < ntiali.ai   1,.  ..■ lie   |ii,.ssiim'   i>   lo   h<>U\   'j I.   il    1" m- 

II ssar\    In  .Aiilaiii   lliis  appaii'iil    iiM-misi-li'iicv  ;  < iiiiivl    ar.'niiiil      I'oi 

a    uicatrr   niiiiilMT   >•]'  i|i>-.iil\  ■■.!    uiiils    lliaii    is    n'pri'sinl.MJ    li>    ll.c   ai-liial 

llllluln'r    ..I     clisM.|\.-.|    IIM.I'Vlll.'s     ^  i     .■  ,     W.'i'.'lll     <<\'    lllssnU.'j    slllist      IIITsl. 

If    was    i,l.srl\,',l    tlial    til.'    pnucr    In    iMillilucI    ti IriMli.-    i-Uircilt        I'llM'trlr 

,.,,ii,|iM-livil\      ill    III.'   c'as.'    i.r   scliitiniis    M.  i;  .    "f   si|..Mi>    wlii.-li    liav.'   an 

nsiimiir   |.i.ssii!c   llial   c.tri     Is  III   llii'   \M  ii;lil    III'  dissdlvnl   siil.sia • 

is  |)iai'lii-all>    hi',   wlicicas        ■•  c |i|.-li\ily  i>\'  tlinsc  snliiliniis  wlii.-li   -Im 

lii.jliiT    iisiimlii-     iHi'ssMic     is  .jMiti'   |ir uiiiTil.      All-lnliiil--    liiailc   llic   ii> 

piillii'sis  liial   III.'  ( ili|rli\i1>    .ii'p.'ii.ls  .III   III.'  spliniiii.'  ..I'  iii.ii.'.'lll.'s  iiil" 

lu,,  ,,i'  1 ■.'  iiniliiiiis  .ir  inns.  .'a.'!.  ..f  ulii.-li  I'aiii.'s  cilji.'i-  a   |...sili\.-  ..r  a 

iH.M^iliv,.  ,  1,,-iii.'  I'liaiu.',  an. I  llial   it   is  ..iiiv    ul sii.-li  .li-.sn.'iali..li  Ul-~ 

thai   III.'  .'!.'. 'lli.'  .'iin.iit   .'an   I ii.lil.'Ic.l  tlir.niuli   tin'  s.iliiti.ni,  tin'  inii- 

s.'i\iiiij  as  il  u.T.'  as  Ij.ials  .'a.ivinv'  lln'  cl.'.'lri.'  .'iliTi'iit .  Wlicii  s.i.liiiiM 
.•lil.iii.li'  is  .liss.iK.'.l  ill  ual.'i-.  it  splits  int..  Na  raii'.v  ill'-'  a  pnsiliv.'  .'Iiai"." 
an. I  CI  rari'Niii'j  ;i  ii."jali\.-  .'Iiaii.'.'.  <>\  N:i  <'l  ,  as  it  is  wi'itt.'ii;  -in  ll- 
..lli.r  liaii.l,  wli.'ii  sii-ai'  is  .|lssnl\..|.  lli.'  iiii.lc.'iil.s  ii'iiiaiii  iinliniKni  aii' 
II. I  I'Ici'tl'i.'  rliaiui's  all'  sil    lli'.', 

Siil.slali.'.'s  ulii.'li  lliils  .liss.i.'iat.'  al'.'  .-all. '.I  . /.  c/n-/.//.  v.  aii.l  tli.is.'  ulii.  ' 
.In   nnl.   Hi, II, U, I  ri'hili'i.      Wlirn    tin'   .'Iri'l  li.-   .'1111. 'til    is   pass,.,!    tlirnimli 

s.iliili f  ili'.-lr.il\  Irs,  till'   imis   wlii.'li   .'aiT.v    a    p.isilixi'   .-liar.,'.,   iims.-   I 

III.'  .'l.'.-li-.i.l.' ni-  p.il.'  Ii.v  w  hirli  III.'  .'iliii'iil  l.'axcs  Ihi'  s..liilinn  llial  is,  i 
till'  saiiic  iliici'linlis  as  til.'  .'iliirnl;  ainl  sin.-.'  lliis  .'li  .-t  i  ml.'  is  .'alli'.l  H 
.•allin.lc,  tli.'s.'  ar.'  .'all. '.I  ..(//.. »v.  Il\  .li-.n^cn  ami  ill.'  iii.tals  lu'lniiu  ' 
tills  '.'iiiiip.  'I'll.'  i.ihs  carrx  iii'j  a  n.'j.'i1  iv .'  I'li.'ir'.'i'  u.i  in  tli.'  .ippnsili'  iliu 
tinii.  auaiiisl  111.'  .'Ill  I. lit  llial  is,  I., war.  Is  III.'  cl.'.-t  r.i.l.'  Ii\  \\  liii'li  Hit'  ."i 
I'l'iit  .III. 'IS,  .ir  till'  aimil.';  lli.'.v  .•im-  I  Ih'I  rlni  c  .'all..!  'Mi...;.v.  'I'licx  in.'lu' 
.1      I,  .1    .,,  , I.I ;.i  ..,■.,,,1,.    vii,.|.  ■.<  ><i  I      I  ■(  >     ,.|,. 

Il    Illllsl    III'  Ull"|i'is|nni|    llial    tills  iliss.i.'ialinll    illtn    Imis    is  ;|  I  l.'ai  I.V    pri'si  1   ' 

ill    til.'   s.iliili.iii    li.'r.iii'   ;iii\    .'l.'.'tii.'    .'iirii'iit    pas^.'s    tlii,ii|i_'li    it.    tlir    mn 


1  I.I  I    11(11      lllMil"    IIVIIK    l>l---i"  I  \'IMl\.    I'>\l/ \l  lilV 


17 


l.riliL'    li..«r\rr    llhil'..lllll.\     iliNtiiUlili,!    .iilclliiliMiit       lliiit     i>.    .•iiliinunl    >'p 

lll.ll    III.'    Il.-J,lliv<'   rlliir'.'.'s   ■■!     Ill''    .■l.lloli^    I Isriv     llrlltliill/.'    ill-'    |M„ili\.' 

I,.ir--'fs  ut'   111.'   iMli..nN       Tin    fi.'.-llii'   .•lin-.'iil   .•iills.s   llii'   rl.rl  |-...|.s   In   I"'- 

,.,,, liMl.jr.l.   Ill |„.sHiv,|\,   III.'  ..tliif   IM'.j.itiv.'lv.  -..   llial   ilh   illlf!)--- 

ti\.'  rc.iv.'  ix  .Mil..!  ..II  III.'  i..ti>  ..r  ..|i|>..>lt.'  M.jn.  'I'lii>  .-nllv.-s  III..  II. "-'il- 
lu.l.v  .'llMl'^..!  l..i.>  1.'  liH.jiiil.'  L.u.ii.U  III.'  |i..xiti\.'  .'I..'!!."!.',  flli.l  til.' 
I..,sili\.'l\  .'liiil-.j.'.!.  l..\»:ir.i'N  III.'  ii.'L-iiliv.'  .•l..-tr...|.'  It  is  tliis  liii-riiil  i.ni 
.,r  III.'  i.iiis  lliiil  .  ii.|.,\'.  s  111.'  s.iliiti..ii  V,  nil  .'..ihiiii'liii'.'  .jualili.'s 

It,  uiil.'i-,  ..r  ill  ,'1  x.iliiti..ii  ..r  :i   ii.'M.  |.'.'li-..l>l.',  III.. 1. '.'111. -s  ..r  II  ,<  •  <.!■  H"ii- 
.  I.'.'l  r..l\  1.'  i\i>t    lliiis; 

II  i>  IM'  IM' 

II  <•  IIH  II  <) 

II. o  II  I  >  II  •• 

III  a  s.pliili.iii  ..I'  an  .L.-li..!;  I-',  tli.'   ih..I.  riil.'>   -|>lil    int..   i..ii>  I'iiin: 

\;i       n       \.i       I  I       \a       »■! 

\,'i        (1        \a        el        \a       CI 
\m       CI        \,i       fl        \a       (1 


Wli.'ii  an  .'l.i'lii.'  I'm'i'.'nl    {.a^-M-  ll i'.:li  a  s..lnli..n  ..!'  an   .L.-l  r.il\  I.', 

III.'   i..ns  li'ii.l   In  anaiiii.'   I  li.in--.'l'.  .'s  lliu-^: 

(\il'i...|i'  .\ii.iil.' 

\:i        \a        \a        fl        n  Cl 

\a        \a        \a       <  1        " 'I  fl 

\a        \a        \a        < '1        •  1  Cl 


i 


1 1  l'..ll.i\\  -,  Ti-.  Ill  III.'  ;il...\ .'  l•■.n■-i.|.'l■ali..ll'^  lliat  //"  i  mitlni'liril  ii  iif  ii  siili- 
sl  iiiii  ill  siihitiiiii  iiill  ill /ll  ii:l  I.  hi  ilii/i'ii  III  irliiili  il  mull  ri/iii  s  ilisxnrni- 
linn.  I'"iii'lli.iiii.iif.  \\'  V..'  a'«sniii.'  Ilial  in  s.i  tar  as  (isiii..li('  prcssUI'C 
I'liiiinniciia  an'  .'.i|i.'.i-ih'i|,  .•.■i.-li  i.  ii  lillla\.^  in  llif  s.-im.'  \\  ji\'  as  :i  iiiiilf- 
.'III.',  llicn  it    l'..ll..\\s  that  tlic  .l.'.-l  li.'al  ilii.'liv  i'\    iinisl  I...  |ii'.i|i.ii'tii>Mal 

Id  ll xt.'lll    In   wllii'll    111.'   ..-llinlii'    |)l-(ssMI'.'   is   L'l.'al.'l'  lllall    \M'  slinllld   fX- 

pri'l   il  1.1  ll.'  rrniii  ill.'  aiiiiiinil   ..I'  sulistaii.'c  ,it'lii.ill\    ilissnlvfil. 

In  the  Determination  of  the  Conductivity  it  is  ..hvinusly  lu'ccssarv  to 
lis.'  slaiiilar. I  I'lin.liti.nis  ..I  .l.'plli  aii.l  \\  i.llli  nl'  ill.'  llni.l  tlii'<>i|._'li  w  liicli  tlic 
I'lii'i'i'iit  is  |),'issi.l,  an. I  III  lia\.'  s.nii.'  standaid  t.t'  ('..iiijiaris.in.  'I'lic  value 
is  til. 'II  knnw  n  as  ///.  spuijii'  ruirliifllril  i/,  tli.'  stall. lar.l  Inr  .'..iiiparis.iii 
ii.-ini;  111.'  .•.in.i\i.'ri\  ii>  ..i'  .i  ii \  |  ..ili.'t i.-.ii  li.|ui.i  v.iii.'li.  \\  .'n.  i..si'.l  in  a 
.'.'III  ini.l.'r  I'lili ',  w.iiil.l  .ilVi'i'  a  f.'sistaii.'c  .il'  I  ..liiii  lictwcrn  tu.i  ii|ip.isit(' 
si.l.'s  .ll'  till'  .'III..'  a.-tiiiii  as  I'li'.'l  ri;i|i  s.      'I'lii'  actual  .l.'l.'iiiiiiiali..ii  is  iisu- 


is 


I   in  SUCH   lllMllM.    I'.ASIS    HI'    I'lIVSKII.IMllC.M.    I'ltOCKSSKS 


,i!'\  iiiiiili'  ill  .-I  cyliliiliii-;!!  vessel  nt'  liiinl  irliiss  (  t'foiii  soft  {lliiss  ciKiUjlli 
;ilkiill  liiiL'lil  I'c  ilissdived  1(1  MlVeel  tile  fesiiltsl.  the  eleetfoiles  l)ei||<r  eirell- 
l.ii'  |ihili  s  III'  |iliiliiiuMi  fill  '\  iMiiieiileij  ill  ;i  l\li(i«ii  ilisljinee  iViini  eneli 
iillnr  111;.  .">  .  *  'I'liis  enihliii't  i\  itv  eell,  iis  it  is  eiilletl,  is  eiilllieeteil 
with    ;i    Nuil.ilile   eleeliie   ;i|i|i,i  rat  Us   fcir   ineasiiiilij;    the    resistance   ii|Te|-eil 


^ 


zip 


l: 

\.j       -        Hi-    .i:     ,  I     I.i.luilv     i.lK       Til.      p'.-itiiiu.i,     ,li-iv     .11,      I,  ,,1,  .,  iil.cl     In      I'll'     lln,  k 

1. 1.1,1      I,,,,-.  IIh  ^       ,',      I..M     111     |...-ili..i-      1. 1      llii,  „  «.illi-,l     tl.i--     lull.-.     llir"n,;ll     win,  !i     llliv     ■"•■ 

.      r..l..i     vvii'i     ll,,      t.  ■  iiiii.ii-     :v      |.l.,;iiii.!ii     Hill'         ir'i.iii     Spin.ii    t 

li\    till-  --olurKii!  til  the  |i:iss;i'_'e  iif  an  ejeetrie  elllTelll    (  Wlieatstmie  r.ri<li;e) 
Me   I 'ill.  (i  .     'I'lie   lesisl.inee   is  of  eiiiirse   inversely    |>rii|iiirt  imial    to   tlie 

i-iini|l|eli\  ilv. 

.\s   .1    s.iiiiH     snliiliuii    is    |ii  o'jressi\(l\    ililiiled.    its   speeide    eiill(lllet  i\  ity 
li.ll  iir.i  ll\     ijei-ie.ises    i.sinee    lliele    ale    now      fewer    liioleellles    lietweeil    ihi' 


ll.'       .         W  :;   .,1   t..,,.       e.iili,,       ;..i      111. 


I l.'t.ll.l. .1,1. 


I  /  .     Ill  I'U.      win; 


Iwii  iilipnvile   l.ii-es  iit    ||ir  eiiit  i  ii  lel  er  eiilie,  aiiil   the  s|);iee   lietween   ions  nr 
inulieiile^    1^    iiicieaseil    .      This    lesiilt    will    lint,    liiiwe\er.    tell    lis    wlietlier 
the  s;ilt   ilvili'  iv  iiiiili'iui.iiii;  any  alteiatinii  in  euiiiliietiii'i'  power  jis  a  eon 
Mi|iieliee,   liir  e\,iiiiple.  nf  .^re.iler  1 1  issneiat  iiili.      To  aseerlaill   this  Ue  Illllst 


!'•■-     '!'•!. '|.,  ,.        1.   1.   IHI'I    .   ,1     II.. I      In      ,!■,.,   1      ■I,..!,, I 1      I. Ill      I.,      .   .|1,   lil.il,.,,,     V,.,|l     I.MllK     ..I. Mill.      I 

I.  'tllii'    till    -I' I   i.il    1, -I-.1.1II.  .-   ..I    .1    v., I, 111. ,11    vxliini-    .-piiih,     ri-l-l.ili.i     is    .1.  riii.t.  Iv    kll..\wi 


i;i,r.(  TKir  coNhn  Tivi'iv.  nissoriATioN'.  Ionization 


l!t 


,  litaii!  liirmcs  n-liitiiiji  1"  Hif  x'hiii  .|Uiiiitit,v  ..f  salt  at  each  .liliition.  II' 
„,■  inultiplv  111.,  spri-ilir  .•nM.hl.Miv  ity  li.V  til.-  vollllilc  of  soliitidi,  ill  <■.'•■ 
uliirl,  ■•,,iitaiMs  I  -raMic.|iiival,.|it  i  s.'.'  |.a-r  "JJ  I .  a  value  will  l"'  >"■'•;"■•;'• 
uliirli  .•.■i.ivs.'lits  the  .■oiiciuclilii;  po^'T  "f  !'  •riain  e,|ui\  aleiit .  Tins  is 
kH„«i,  as  I  he  ,,iinn,l,  „t  or  n,oh,i,hir  ro„.h>,-l,r,l  „*  an.!  is  represeiite,!  l.v 
111,,  simi  A.  When  il  is  <letermiM...l  loi-  proj-ressively  .iilute.l  solutions. 
A  uraihially  iileieas..s,  iinlieatiii-  that   //-.   <  jjiri>  „ril  «-/  //"   ,l,rln>ll,t<   //.sW/ 

„s  „  nnulmlnr  nnn„^,s  with  .HI ,il ion.  1 ails,,   it   -lissoeiati's  iiii.re.     Tiie 

,.M,.nt  of  tliis  iiier,.as,.  is  fouii.l  to  lieeoiii,-  less  an. I  less  as  .lililthu. 
p,„...'...ls.      I?>    pL.ttiii-  the  vahi..s  ..r  the  iu.,l..eiila|.  e.m.ilH-tivit y  of  sil.'- 

.■,s>iv..  .liliitions  as  a  eiirv...  tli.'  vain.'  at  iiitiiiit.-  .liliiti -an  1...  as....|.taiii...l 

hy  ..\llapolati..ii.     This  valii..  is  r..p|...s..nte<l  l>y  A^^. 

N.,\v,  l..t  us  s..e  h..\v  tlii.s..  facts  i...ai.  ..lit  th."  tli....ry  ..f  ..l....ti...lytic  .liss.M-ia- 

li.,i,.     A.-cnliii-  t..  this  hypoth.'sis  th ii.lii..tivily  .l..p..n.ls  ..ii  th,.  iiiim- 

Imt  ..f  i.iiis   IS....  pa::..   17'.  an. I  sin.-.,  it   is  at   a   iiiaxiiiiuiii  at    iiitinitc  dilu- 
liiili.  Ill,   fiiliK  A  .    ii'iist  rii>ns,iil  lln    lnt,il  iiiniih,,- of  ions  Ihul  niii  hr  prtt 
,hi,,,l  hii  llu   ,lis>ori'itioii  of   I   ,i,;n,i-,ii,nr„l,„l.  mol  A  Ihul   "I   so,,,,    olh<  r 
,l,l„lo,„.     If,  tli..|..-f..ie,  w..  .livi.le  A  l.y  A  ■     ".'  .ihtaiii   a   valll..  i.^all...!  ..^ 
«hich  must  |...pr(.s..nt  the.h.irn...  t..  wlih-h  th..  el....t|.olyt..  is  ioiii/...!  at  th.. 
various  .liliiti.uis  at   wlii.-h  A  is  nieasure.l.     Kr..iii  "hut  Ims  l...eii  sai.l   c 
..'ar.linir  th.'  ..smotic  pi-.'ssiii..  of  similar  solutions,  it   is  .'viilenl   that   th.. 
valu..  a  .-..ul.l  als..  he  eal-'ulat...!  l.y  timlini;-  the  ..xteiit   t..  which   the  .l.'- 
p|...ssi..ii  of  fm./iusr  point  A  is  .^n.ater  than  w.Mihl  1...  espi'd...!  fn.m  th.. 
niiiiilM.r  .,f  .lissolve.l  mol,.ciilcs.     As  a   iiiatt.-r  of  fuct.  it  has  l.e..n   toiiml 
Ihat   practically   hhiitical  valil..s  ar..  ..htain.'.l   for  mnny  sulistaiic..s,  thus 
fiiniishin..'  almost   iii....iitrovertil.l..   pro.'f   iu   supp.iit    of  the   .liss...iatioii 
hypothi.sis.     In  th..  cas..s  of  u.-al<  a.-i.ls  aii.l  hasi's.  it   is  iiossil.l,.  t.i  secure 
a   value,  call...!  the  iHssorinHoii  roiislmil    iK>.  \\hi..li   repres.'nts  the   rela 
live  values  of  „  at   all  .lilulions.     Sin.-c  the  a.-livity  of  a.-i.ls  ami   l.ases 
i-,  .l.p.'n.h.nt  iiii.iii  till-  iiumher  of  11     aii.l  (»ll    ions,  respect i v.. ly.  si.f   fiv 
liy  (lissociati.in,  it   foll.iws  that   it   must  li.-  i>r.>p.uli.>nnl  to  K,     Il   will  lie 

n ssary   t.i  postp.me  ;i  consi.l.. ration  .if  the  applh-athui  of  this  ... instant 

until  \v.-  Iia\.'  stu.li.'il  mass  action  i  pa'_'..  '_':!). 

Biological  Applications.  Th..  luai'tical  value  of  sii.h  kno\vl...l'_'..  r.-sts, 
not  s.i  mii.'li  on  aii.\  .lir.'.-t  simple  .-ipplh-ation  that  can  \h-  mad..  .>f  it  in 
explainiiifr  ))liysioloi;ii'  pr.i.'.'sst.s.  as  .in  the  (.ss.'iitially  imp.iitaiit  li..arinv: 
\\lii..|i  it  has  in  ..nalilint:  us  t.i  iiinlersian.l  th.-  iiatur..  aii.l  .ip. .ration  <if 
.illu'r  physic. ich'.iiiic.-il  fa..|.  -s  ...ini'criie.l  in  jih.v  siol.eji..  iinu'i'sses.  With 
out  Ji  ..h.ar  c.imprt.h..nsioii  of  the  ..1.. mental  laws  of  .iiss.iciatioii,  it  is 
inipossilil..  I.I  consider  sii.'h  |irolileiiis  as  tlios..  whi.-li  cone.. in  the  a.-tivitics 


■Im   Mill,  I    WMi.U,   111,-   'iHil,,  iil.u    ,',ii.|.i.  Inilv    i~   .1:,    ^i.,,ilu 
IkT.imi  ,niii\al<  lit'  I'liit.iiiu '1  ill    1   »■  ," 


it>    ,li\i'U-l   liv   tin    lunni,!  r 


•J(l 


I'llVsicnciiilMUM.   liAsis  OK   I'll Vsiul.oiili '.vr.   l'|{(iti;ssi;s 


of  ('llZ,\liirs  (  lil.lNS  iiclidli,  ell'.  I.  Illi'  nri-iiirfli if  flcrtric  (MIITcllls  illllillU' 

till'  iilivsiiiliiuic  ii(li\it.\  "f  iiiiisclcN,  iil.-uiils,  ;iiicl  iiciVfs.  :im<I  llif  all 
iiiipiiflMiit  i|iii'stiiiii  111'  tlif  rc.icticiii  m-  II  inn  (•niiciMil  ral  inii  ol'  tin-  Imilx 
llili.ls. 

I'.cfiiii'  |)icicici|iiin'  til  sliiiw  liiiw  tlii'sc  farts  fipiifi'iiiiiii,'  tlif  nature  nf 
Milutiniis  air  aiiplicalili'  to  llu'  stinlv  ol'  |)liysiol"i:ic  proci'SM's.  it  may  In- 
well  (o  iiiiliratr  oiii'  or  two  iiislain-i's  in  wliirli  nimsiin  iiii  ii/s  of  iliitnnil 
ciiiiiliii'llrihi  mill  III  ilissiii-iiiliiiii  Iniri  iJinrl  }ihtlsiiihiil'H-  nillii.  Tilt'  cilVll- 
latioii  liiMi'  of  till'  liloodtlou  tliroimli  an  oriraii  «'aii  Ix'  (Ictcriniiicil  liy  lir.st 
tiinliiii;  llir  I'lci't lii-al  ii'sistaiici'  of  a  slimt  pii'i'c  of  tlii'  vein  of  tlic  orj;an. 
ami  llii'ii  oIiMTxiiii;  llii'  ciiaiiu'i'  in  rcsistaiu'c  wliii-ii  is  inoilm-cd  w  lu'ii  tlii' 
coiiiiniiiv  ity  of  till'  lilooil  in  llic  vein  is  .'ilti'i'ril  li.\  tlic  arrival  in  it  of 
salini'  iiiji'i'ti'ij  into  tin'  ai-lri-,\.  Tiic  intcrxal  clapsinu'  ln'tw  ci'ii  tin'  iiiji'i- 
lion  into  tlir  aitciy  am!  tin'  <'lian!.'i's  in  rcsistaiu'c  in  tlic  M'in  olivioiisly 
('i|nals  till' ciirnlatiiin  tiim'  iH.  \.  Sti'wait). 

Till'  saiiH'  in\  I'sti'jator  lias  nscil  iiicasiiii'iiii'nis  hy  cli'i'tiii-al  romliii'tiv- 
ity  to  study  llir  passam'  of  rli'i-t  i-olyli's  out  of  tin'  ri'il  lilood  corimsidcs  into 
till'  siriii  :.  Inilcr  mniiial  romlitioiis  tlic  l:lood  scruiii  lias  a  ccrtaiii  I'li'.' 
tiiral  i'omlui-li\  ity  I'lpial  to  thai  of  a  1 1. II  pi'i-  i-i'iit  sodiuiiii'lilinidf  solution. 
Till'  i-ondiicti\  ity  of  tlu'  dililninati'd  Mood  is  only  aliout  onclialf  that  of 

si'inni.  I aiisc  it  rontaiiis  i'oi))usrh's  w  liicli  aii'  iioncondm'loi's  and  tlicri' 

fore  olistrm-t  thi'  free  jiassau'"'  of  tin'  ions,   just  as  a  suspension  of  (piait/ 

poudiT  in  a  sodium  chloiidi'  soliilion  lowois  tin iidui-ti\ity  of  the  hit 

tir.  If  aiivthinir  oi'eiiis  theict'me  to  oeeasimi  a  [lassaire  of  the  saline  eon 
ti'iits  of  the  coipiisi'hs  throu>;h  their  walls  into  the  seiiiiii,  an  increase  in 

thi'  clei'tlii lidurti\ity   will   he   pioillleed.     Tile  value  of  this  method   in 

the  iii\  e^tiuatioii  of  I'hanaes  in  /n  nii< nhililji  of  the  red  eori>llseles  is  de 
pendent  on  the  faet  that  sm-h  mii:ration  of  electrolytes  out  of  tlie  eor 
piiscles  may  occur  licfoie  any  of  tlic  less  dilTilsilile  heiiioiilolpin  itself  has 
csi-apid.     The  rise  in  cniiduclix  ity  |ir dcs  the  hcmolvsis  (see  pas;e  7). 

AlthoUL'h  dclci  l■lillation^  of  the  spccilic  comliiclixity  of  hlood  and  tiriiie 
under   \aiious    pathologic   conditions   have   also    lieeii    made,   the    result-- 
liaxc  not   liecn   found   to   possess  aii>    diairnostic  value  or  clinical   siiinili 
i-ancc       .Measiiremcnts   of  llie  elediic   conductivity  of  Mood    have,   hou 
e\fr,  lieeii   used  1p>    Wilson"  and  Ip\    I'riestley  and   llaldane"  to  detect   tie 
deyn f  dilution   when    lai'.'c  ipiantilics  of  water  are   inc-ested. 

Another  application  of  cniidiicl i\  it \    measnremeiits  in  liiocheiiiist ry  ha- 
liceli   made  in  studyilie-  the  diei'slive  .-letion  of  proteolytic  enzymes   (  Ma\ 
lissi.     The  ueiieral  action  of  the  enz\  ines  is  to  Ineak  the  laru'e  undissn 
elated    nioleciiles    of    tiie    higher    proteins    lalliuiniii,    casein.    ete.K    int" 
siiiidler  molecules  uimiiio  acids,  etc   ,  w  hieli  ;ire  pailly  ioiii/cd.     .\s  iliecN 


i:r,i:(Ti{i<'  ((iNhiiriviTV,  nissdciATiow  ikm/.atiun 


!1 


liiiM  piccccds,  llicicl'iiii',  llic  cmiiluctivity  of  tin-  diucsl'mii  iiiixliiti'  piu- 
..Mi'ssivi'lx    iiicic;is('s.  iiii'l   is  II   MU'iiMiif  i<\'  tlic   Tiilc  iil"  (liin'stic)ii. 

Ai>i>llciili<>iis  III   llii   ilissiii-iiiliiiii  liiii>i>llii.sis  ill   piivsiiilniry  (Mil ni  tin- 

c  \|.liiii;iti r  Mii'ii    iilicimiiiciiii   MS   tlic   piniliicliiin   iit'  I'lictric   iMiri-ciits 

(liiiiiiy;  iihisciiImi'.  lilMinliihii-.  ;iinl  mitmius  iii'li\it.\.     'I'l \iii-t   <|ct;iils  nC 

llii'  .•i|i|)lii';ili(iii  lire  imt  ns  vet  MinicicMt iy  iiiKlfislnnil  to  \\;in-;iMt  <>iir  ;it- 
Iriii|>tiii!,'  til  (In  iiiiHf  lliiiii  iiiiiiciitf  tlic  irc'iicnil  lines  iiloiiy:  wliidi  the 
piiilpli'iiis  ;irc  liciiiL'  iiivi-stiyiitf.l.  I.ct  lis.  for  cxiiiiiidi',  cniisidcr  Imw  tin- 
ciirii'iit  lit'  ;ii-tiiiii  111'  Hillside  liiiiy  lie  expliiined  ill  terms  uf  tlie  ilissnejiit imi 
liypiitliesis.  Til  (III  sii  we  must  delve  il  little  flirttier  into  pliysieoelleiii 
ie.il   leseiiieli,  wlieii   we  slijiU   find  tliat   there  ;iie  two   ruftlier   t'nets  euii- 

eeriiinsr  iiiiii/ed  miileeiiles  tluit  must  lie  uf  im|iiirt;iiiee  in  einii timi  with 

nlir  priilileiii.  The  first  is  that  the  emit rilmt inn  wliieh  eaeh  inn  makes  tn 
the  eipiivalellt  (iir  ninleeillari  eiiMillietivity  uf  a  snlutioll  is  iliilelielldeiit 
iif  llie  other  iiiii  with  whieh  it  is  associated:  and  the  seeoiid,  that  ions 
ditVer  coiisideralily  in  their  ciiliilint  ini;  power.  Since  the  iinivaleiit  ion-, 
K..  Nil..  ('!/,  No  ',  carry  ehaiLTcsof  the  same  mairnitude,*  and  yet  all  dn 
mil  coiiiliiet  to  the  same  dc"_'rec.  they  must  iiiom'  at  dilTcreiit  \elocitic-i 
throiiu'li  the  solution.  We  are  driven,  tlurcfore,  'o  the  conelusioii  that, 
expiised  to  the  same  eU'ctric  fnree,  ililTcicnt  imis  liave  ililTcreiit  iiioliili- 
ties;  that  is  to  sa\ .  when  an  electric  eiirrciit  |>asM's  throuirli  a  solution  of 
an  electrolyte,  tiie  positively  ehaiircd  inns  move  touards  the  cathiulc  at  a 
dilTerent  rate  from  that  at  which  the  iieeatively  cliari,'cil  ions  move 
towards  the  anode.  ( 'oiifirmation  of  this  eoiielusion  is  olitained  liy  exaiii- 
iiiation  of  the  eoneentrMtioii  chaiiiics  aroiind  the  two  electrodes  of  an 
electriil\  tic  cell.  The  actual  velocity  of  eaidi  inn  can  lie  determined  liy 
ex]>eriiiieiital  means. 

•Tliii*    I'.ir.ilnv   »li..«..l   lli.it   tlic   :itTi'.iuit-  nf   tin-   \;irf'ii>   i"M>   lilnr.il.d   liy   <  In  tmlv -i"   :irc    in   the 
^.iiiir   i^iti"  .1^  tlit-ll-  iliin.i.il   I  ■imw.li  iit^. 


CIlAlTKi;   IV 

TiiK  i'i;iN<  ii'i,i;s  iNVoiAiih  i\  TiiK  i>r/n;i{MiNAi'i(tN  <»i-  Tin: 

IIVhUoCKN  ION   ('(iNCKNTU'ATloN 

TITRABLE  ACIDITY  AND  ALKALINITY 

All   iiri.U   li;i\i'   niic    |>iu|iri-|v    ill    1 iiiiiiii     iiaiMcIv,   tluit    llicy   iM.iitiilii 

lixdnii^cii     aiiil    ulini    tlic   afiii    Imm lo    nciil  lali/rd,    it    is    tliis   .•Iciiu'ii' 

wiiii-li    IpccniiirN    icpla I    li>    M.I illiiT   catinii.      Kv  i.Iciitl\ ,    llicii.    tlir 

stri'imtli  lit'  an  arid  is  piiipm  liniial  to  llif  luiiiiln'i'  id'  displac-calilc  lixdm 
>_'cii  atdius  'Ahirli  it  iMiiilains.  It  may  vinitaiii  utlicr  iiydni-rcii  atmii-- 
\\liiidi  ail'  so  lioiind  up  in  llir  iiinh-.-iilc  tiial  llif\  do  iidI  lifconic  displaced 
i\hcii  an  alkali  is  iniM'd  with  tli.'  ai-id.  I'm'  cxaiiipl'',  in  orLMiiic  arid - 
likf   ai'tlic,   <'II('(M>|I.    it    is   nnl\    llic    II    atom    attaidicd    tn    lln'   ('(><»ll 

L'n.iip,  lint   lint   tliu-c  attaclicd  \u  tin'  fll     umnp.  that   is  rcpl; aid.'.     It 

must  tlirnrinr  In-  pnssililr  t.i  |ircpaic  Tor  (■\in\  acid  a  sidiitimi  liavinv 
exactly  III.'  Villi, (•  nciitrali/iii'j;  pi.uei-  as  tliat  i>\'  aii.\  ntlier  acid:  that  is 
the  same  vuhiine  i>\'  sululinii  must  lie  icipiiied  in  eacli  case  tci  neiitrali/' 
a  '_'iveii  i|uantil,\    nf  .-dkali.  the  p'>int   ni'  iieiitrali/atioii   lieiiiu:  ju'dired   li,\ 

the  clianuc  ii Inr  n\   indicatms.     As  a  vtaiidaid  a  'iiaiii molecular  snln 

tiini  nt'  an  acid  with  one  displai-ealile  II  ion,  such  as  liydrochl"i-ic.  i 
cdioM'ii.  This  we  call  a  '•iicn-nial  acid"  iN  .  To  prepare  a  iioinial  soiu 
ti<in  o|"  MciiN  lia\  iii'u  tuo  dJNplacealile  II  atoms,  -.uch  as  ll.Stt,.  we  can  im! 
ho\\e\er  use  a  '^laiii  iiioleciilai-  ipiantity,  Init  must  take  niie  half  of  il 
Mini  siinilarl.N  in  the  ca-r  ,,(  those  with  three  II  atoms,  such  as  1 1  I't  I 
a  one  third  'jiaiii  molei'iilar  Milntioii  will  he  a  iiorm.d  acid  solution.  I''" 
practical  piirposrv.  use  is  vmy  ',;enerall\  made  of  solutions  that  are  soii! 
fraction  of  the  normal,  c.  l;.,  tenth  or  deciiiormal  i  written  N  H'  i.  or  Inn 
dredth  or  cciitiiioinial  <  N    Mm).. 

In  a  similar  wa\,  alkaline  solutions  c;in   he   prepared,  a   normal   alka 
jiriiiv'   one    which   e\actl\'   corresponds    in    strenutli    with    a    normal    aci 
(i.e.,   can   exactly   neiitrali/.e   it    .      Now.   the   eliaraeterist  ie   of  alkalies   '■ 
that  they  produce  in  solution  "<  Ml  "  m-  liyi|rox\  1  ions;  so  that  the  proi-e. 
of  neutiali/alioii  must  consist   in  the  union  of  the  II  ions  of  the  acid  w  i' 
llie  (HI    ioiis  of  the  .alkali   to   form   waier:   K<>ll      IKI         KCl     II  (t.      W 
c;iii,   thcrel'orc.   pict'are   normal    solutions  of  alkalies   liy    dissohinu'   in 
liter  of  water  such  cpiantitics  of  alkali   i  in   L'laiiis^    as  will   \  ield   the  <HI 
reiiuircd  to  I'cact   with  the  a\ailalilc  h\ilroeen   in   uoriiial  acid  sojutinii 


IIVl>llO(li;\  I(l\    CONCI.N'lltATHIN 


2:1 


Actual  Degree  of  Acidity  or  Alkalinity.  Airnnliny  i'.  lli''  I'.nv.z.'in-.' 
nirllKMl  ..f  litnitiiMi  !i  iioiiiiiil  solution  of  a  |.o\v.-ilul  tniM.iiil  .i.-i.l.  su.-h 
iis  liydiocliloiic.  is  no  slimmer  tliiin  ii  noiiuiil  solution  of  :i   u.'iiU  jo-id, 

surli  ;is  ii.Tlic  or  Inctir.     It  rc.|nirrs  no  f.-wcr  .•.<•.  of  .V  iilk.ili  to  1 trnli/r 

it.  I?iit  the  nornnil  solution  of  tin-  poucrful  iici<l  tiistrs  ni.nc  ;n-i,|.  is 
more  toNie,  .lissolvcs  nieliiis  more  rcnlily,  iiinl  in  iill  its  other  eliemie:il 
■.\ui\  l)livsiolo-.'ie  properti.'s  iiets  niueli   more  c|niekl>    tli.in   tli.'   ueaU   Meid. 

>o  tliat  llie  lilriihl,   ,i,-i,filfl  or  ,ill,;ili,nl  11  r: I  express  tlie  real  slren-tli 

of  tiie  aei.l  or  alkali,  or  tlie  aelnal  de-r f  aei.lity  or  alkalinil.x .      It    is 

in  this  eonneetion  tlnit  the  .lissoeiati..li  livpoth.'sis  aids  us.  foi-  il  sn--esls 
that  the  de-jree  to  whieh  the  aeid  I.eeomes  dissoeiated  into  11'  an<l  the 
ivinainder  of  the  moleeuh'  will  determine  its  real  strength  1  see  paiie  IC.I. 
The  .ph'stion  is.  h..u-  arc  we  to  measure  the  latter.'  (»iie  aetlnn  of  II  huis 
whieh  we  may  nu'asure  is  that  kin.wn  as  r;l<ihiliv  that  is.  the  power  to 
aeeelerate  reaeti.nis.  sueli  as  the  splittiiii;  of  c-ane  su-ar  ( (",,ll  ^  <  >,,  1    ml" 

!.dneose  and   le\  idose,  whieh   otherwise   would    pro( 1   v.'r\    slowly    (see 

pa^e  7:.).  If  then  the  real  stren-.'th  of  an  aeid  depends  on  the  deL'iee 
of   dissoeiation    whieh    it    uinler-jocs,    tij-ures    repres.'ntintr    the    eatalytie 

power  should  eorrespoinl  with  those  re]*,. ■sent in-,'  the  relative  c luetivi 

lies  of  the  aei.ls  in  e«pii\alent  eoneeiit  lation  1  s.m^  pa-re  I!»i.  That  this  is 
a.'tiially  the  ease  is  shown  in  the  followin-jr  talil.'.  in  whieh  the  ahove  values 
of  various  acids   are   jriveii   eompaivd    with    IK  1.    whieh    is   taken    as    KXt. 




.\<'ll> 

CAT.M.VTie  roWKI! 

i;ki 

.vrivi 

eoNhi 

■TIVITV 

1(111 

ill'] 

MO 

|)i.'lil..ii„'elir 

''7 

-■' 

MiiiiimIiIo1!iic 

ic                                l.s 

l.'.i 
1.7 

Foiniic 

1..-. 

.\i-rli.- 

(I.IO 



(i.  IJ 

It   will  he  evident  that,  if  we  eouhl  measure  the  eoneelitration  of  fi 

II  ions  in  a  solulioii     that  is.  of  II  ions  that    ■•••  not  matehed  hy  Oil   ions  ~ 
we  should  have  a   faithful   index  of  its  real        idity.     This   measurement 

has   heeii    rendered    possihie   liy    the   aiiplieati f   two    other    physieo- 

eheiuieal  i>rineiples  naiiiel.v.  those  of  mass  action  and  eleetroiiiotive 
force.  Sim'c  the  ohji'et  of  this  xolunn'  is  to  present  the  seientitie  hasis 
for  the  various  iiielhods  that  are  used  in  iiioilern  iiiedieiue.  it  will  he  nei- 
essary  for  us  to  revii'W  the  iiiaiii  principles  of  tliese  two  actions.  We  shall 
see  that  they  apiily.  not  onlv  in  thi-  measiirement  of  II  ion  concentration, 
hut   in   many  other  physioloeic  processes. 

Mass  Action 

When   materials   taki'   )>art    in    a    reaction,   some    molecules   are    ih'ccnn- 
posiii};   while  others  arc   heint:   fdrmetl.      After   some   time,   however,   a 


■2\ 


I'llVMi  IK  iiiMir  M.   ii\si>  (ir  I'livsKii.iMiK  \i,  cKiHi;; 


ciiiHliliiHi  is  ii'Mi'liiil  ill  wliii'li  ilic  i-liiiiiiics  ill  mil'  ilirci'tiiin  .'iri-  cxmiMIx 
olTsfl  liy  tliiiM'  ill  llic  iitiirr.  All  ii|ilililiriiliM  i>  sjiiil  to  li.'ivc  lici-iniic  CNtilli- 
lislird  iirl  \v  ci'h   tin'   ii'jicl  iiiir  Milist.iiicc.s,      ruMiiiiL!:  in    miinl  lii.it    llic   imis 

.•linl  lllolrclllc's  I'lltirilli;  illhi  llloc  lr;|c-li(ill>  Jirc  i"  ill>t  Jl  lit  I V  ll|ci\illLr  illiinit 
illiil  ciiiiiiim  ill  c-ii|lt;irt  with  niic  jillntlii'l-.  it  i>  c.-lsy  tii  m'c  tlliil  it'  w  !■  WiTC 
to  mill  .'III  :ii|ilitiiiii.'il  i|iiMiitit\  lit'  mil'  Iviml  nl'  iiiult'i-iilc  nr  inn,  tlicrc  wmilil 
111"  ii  fii;iii!.'r  .'ill  .'iIiiiil;  tlu'  liiii'  until  'i  iiru  i'i|iiililii'iiini  wjis  I'st.'ililislii'il 
It',  nil  till'  ollli'l'  llllliil.  \M'  well-  tn  I'lliiiiM'  iilir  Mini  of  llliilliMlIc  iil'  inll 
.•IS  f.'ist  .'IS  it  is  t'lil'liiril.  tlir  ri{liililil'iiiiii  riilllil  lif\  I'l'  In'  rstjililislii'il,  .'iliij 
till'    n'.'irtiiiii    wiiiiiil    prni'iTil    until   nil   nl'  lliis   in.'iti'i'i.'il    liini   ilisiippcjircil. 

'I'lii'  iiatiiriil   I'iiti'  iit   wliii'li  iiiiy  I'lu'iiiiriil   I'l-iii't  imi   |ii' 'I'lls  is  drpciiiii'iit 

npnii  ii  niiiiilii'i'  111'  I'linilitiniis.  siirli  MS  I'lii'iiiii-iil  iitliiiity.  ti'iiiprfalufi'. 
cjit.'ilysis.  ami  rmii'i'iil  nitimi,  (  M'  tlirsi'  i-niiilitimis  lli.'it  m'  I'nni'i'iitr.'itinii 
is  ninst  ri'iiililv  llli'.'lsllli'il.  ii'  \M'  IM.lilltaill  ;iil  nl'  tlir  I'nllilitinlis  iitlirl' 
tinill    tll.'lt    III'  rniii'i'litl'iltinll    lllli'li.'lliuril.    .'linl    iji'siiru.'lti-    tills   i-nliiliilli'il    ill- 

lliii'in'i>  .Is  K   ii'iiiisiaiit    ,  \M'  sliall   liiiil  tll.'lt   till'  sp I  111'  till'  ri'ai'tiiin  is 

plnpnltinll.'ll     tn    till'     lllnli'iMI  la  1'    rnlH'l'll  I  I'al  inll     n  l'    till'     ll'ai't  i  II  y:    Slllist  aill'CS 

I  i.  I'.,  the  iiiiiiilii'i'  nt'  LM'aiii-iiinli'fiilai'  wi'i'jiits  piT  li'i'i'i.  In  ntlii'i-  wm'ijs, 
till'  spiTil  with  w  li  li  \\\>i  Mllistaiii'i's.  a  .'iliil  ii.  Miitr  tn  I'nriil  ntlli'l'  sllli- 
st.'IIII'l'S.    I-    .'lIliI    il.    «il|    III'   i'\pllsv,(|    |i\     ll lllatinll, 

I,        :i  i  1..     r--    k'       .-,    .    .,1  ,  ;■ 

wliirli  iiii'.iiis  that,  wlirii  till'  i-rai'timi  is  i-nnipli'li',  tin'  i-niiipusit  inn  nt' 
till'  iniMiiir  will  III'  ili'priiili'iit  iipnii  till'  ratin  lii'txM'iii  k  ami  k'.  Sinrc 
linUiM'i'  tl:i'>i'  air  hntli   rniistaiit'.,  thrir  i|Uiitiriit    is  alsn  I'mislaiit    (l\i.  anil 

^1, 


\\i'    h.'iM'    till'    ripiat  inn. 


i| 


K.    imlii'.'itiii'^-    lliat    111!    iiiatti'r    linw 


tlir  rnm-i'iitratiniis  a,  li.  r,  ami  i|  .'iir  \ai'iri|,  rrai'tinii  will  t.'ikr  plai'i'  in 
niii'  iliiii'limi  111'  till'  iithir  until  tin'  I'niirriit  rat  imis  lime  ln'i'iiiiic  ailjlistcil 
sii  t  hat    K    li'iiiailis  nii;'|i.i;M^ri|. 

As  an  r\aiiiplr  nf  tin'  applirat  inn  nl'  tlirsi'  laws.  Ii't  ns  taki'  tlii'  rrai'tinri 
whii'li  iic'i'iiis  lirt  w  I'l'ii  ah'iilinls  ,",ml  nruaiiir  ai'iils  tn  I'nrni  thi'  siilislaiii'c; 
ralli-il  i-.t,Ts  a  rrai'tinh  whii-li  !■>  aiialii'_'nns  tn  that  lii'twi-i'ii  iiiiiii'i'al 
.'ilkalii's  ami  ariiK  tn  I'nrin  miit  ral  s;iltv,  .'hkI  wliii-h  is  nl'  sprrial  iiiti'i'i'st 
In  ns  lii'i-;iiisr  it    i>  thr  rrai'timi   iii\ni\i'.|  in  till'  >plittin'_'  nt'  anini.'il   I'afs. 

Till'    ripi.'llinll    i'lir    till      li'ai't  inn    is; 

I'    II    I  III        I'll    I'lKlll     .-III   Ulll'lll  II   I). 

..  Uivl  :..,'i;.  I.  II,^I    :  ,,  t:,t.  . 

:il."l.'.l  ;,o,|  .  ;,ii,-l,l) 

( >!'   cxpiTv-i'il    .'iri'niiiiii'j    In    i|i,'    hiw    ill'   iiia>s    ai'tinii: 
|i'  II  I  111  i      ;  III  I  I II  III  I 


•Ih. 


.k.I-  llhll,  , 


jl'JI  <MII  I'll    I       I  ||,()| 
il.ii    .'....'.    i;  . .!..■,',,    .,..,.,,,1111, - 


k 


IIVI>l((«il;N-H>N    niNcllNTUATKlN  J-> 

Now  it  is  cli'iii'  tliiit  it'  \\i'  iiicrciisc.  sii\ .  II J  >  in  tin'  iilicivc  i'i|iiiiti<)ii,  tln'ii 
ill  (irdi'i'  tliiit   K   iiiiiv-   I'liiiiiiii  iiiicliiiM;.'i(l  CJI  OOCt'II,  mnst   diiiiiiiisli  nr 

the  siilistiMici's  wliicli    I'oi-iil  till-  iiiiiiKTiitDr  of  li i|iiatiiiii   iiiiisl    iiii-i'iMsi'. 

.  !•  Iiotli  tlicsc  cliaii^ri's  iiinst  diTiir.     As  a  iiwittcr  nf  \';\rU  in  sni'li  ii  case  as 
liic  almvc.   liiilli   of  these  ad  iusliiieiits  taki-   lilaee.    fur.   as   llic   ester   lireaks 


down,    it    must    tliereliv    inerease   tl 


le    eoiiei 


iiti'atioii    of    acid    and    aleoli 


Siiiee  in  aijiieoils  solutions  the  I'eaetioii  oeelirs  in  tlie  presein'e  of  an  exeess 
of  water,  it  is  cviih'iit  tliat  the  teiidcney  for  an  ester  in  the  presence  of 
water  is  to  lireak  down  into  aleoliol  and  aeiil.  and  this  niiist  oeeiir  in  all 
leaet  ions  in  tiie  liodv  tlniijs  in  xviiieii  wali'i-  enters  into  tlie  ciiuMtion. 

Physiologic  Applications.  The  apiilieation  of  the  law  of  mass  ai'tion 
ill  tlie  explanation  of  liioelleiiiieal  pi-oeesses  is  \eiy  eNtensi\«".  .Most  of 
the  rcnctioiis  \\liieli  eii/yiiies  or  ferments  are  eapalile  of  intlneiieiii!,'  are 
of  the  same  jieneral  nature  a  ;  that  represented  alio\c,  and  ihe  jiroduets 
oT  their  aeti\ities  are  usually  the  suhstanees  on  the  side  of  the  t'i|uatioii 
ill  wliieh  no  water  molecules  appear  i.e.,  tlii'.v  arc  liydrolytie  reactions. 
l".ii/\  iiies  merely  accelerate  the  reaction  (  pauc  ~'l  > .  so  that  if  we  start 
uilli  a  mixture  of  the  sul)stan<M>s  on  either  sidi  of  thi'  ('(pialion.  all  they 
ilo  is  to  ;iccclcrale  the  prodiic-tion  of  a  sullicieiit  concentration  of  those 
on  the  other  side,  until  the  c(|nililiiium  point  is  reached,  j-'or  example. 
I'll  enz\  me  present  in  pancri'atic  juice,  calleil  ///)</.<<,  accelerates  the 
liicaUdow  11  of  such  esters  as  neutral  fat.  wliiidi  consists  of  tlic  triatomic 
;il  ohol.  ulycerol.  comliined  with  the  fatt\-  acids  palmitic  ((',  "  ,'  'X*'!', 
steari<-  If, -ir,  cool!  ■   aiol  oleic  i  ( '.11    ( '()( »ll  i : 

<',II.  (O  or  "•,11    I      ::ii  (lii:^  ;;c,  II  rooii  cif    lOll  i  . 

(tlie     ni'litl.'il     ImI,  (  llie  r.lllv  Mi-i.l.       Iu;l\rrl.i|  I 

ti'i-ti'inin  1  sli;iiii-  i 


I  I  ili'r  ordinary  conditions  tin-  reaction   proceeds  until   nearly  nil   the 

iiiu!i,i!    U[\    has    liecome   di mposcd    Iiccanse    of   the    pre|ionderaiice    of 

water,  hin  if  we  start  with  a  iiiixtnre  id'  fatty  acid  and  irlycerol  \vith 
.iiist  enninri,  water  to  permit  tiie  en/..\  nii'  to  act,  the  reai'tion  will  pi'o- 
cicd  in  the  opposite  direction  i.e..  so  that  some  neutral  fat  will  he 
^."  III  liisi/i  il.     This  is  called  the  n  fi  islhli   miiuii  nf  iii.iiiHts. 

localise  of  the  uiiivcrsjil  presence  of  water,  it  is  jilaiii  that  sucli  re- 
'isiMe  reactions  could  not  alone  he  lielil  respoiisiMc  for  the  synthe- 
^is  of  iiciiiijil  fat  or  of  similai'  sidi>-taiic-es  in  the  animal  Imdy.  The  only 
V  ay  li\  which  synthesis  could  oci-iir  under  these  conditions  would  he 
ii  the  siihslaiice  producccj  aloip^  with  the  wjitcr  wcii'  removi'il  from  the 
■-  !'■  of  the  reaction  as  soon  as  it  was  formed.  This  miiilit  occur  hy  the 
I'll  cipitatiou  of  the  sulistance  or  liy  its  heeomiim  surrounded  l>y  an  en- 
■ilnpe   of  some   inert    tnaterial.      In   the   sxnthesis   of   neutral    fat    which 


2(5  niYMr.lClllMMAI.    IIXSIS    UK    Cll  VSlCKMltr  \l,    I'UlM  TSSLS 

,M.,'Uis  in  ll...  ..|.illM'liu.M  of  til..  ihlrsMi..-  out  u(  tlw  fatty  i.ri.l  ni..!  -lycTol 
al.s..rl..'.l   from  Uu-   i.il.stinal  n.nfnts.  it   is  ,.osm1.Ic  tl.at   tl..-  last   in.-ir 

,i.,„r.l    rro.Mss   IMS.      lluT   rasos  tlu'   sul.slaur..    may    l.c   .•air.r.l 

iiway  l.y  till'  Moo.l  or  l>m|.li  or  urine  as  fast  as  it  is  f..rMi.Ml. 

The  Law  of  Mass  Action  as  Applied  to  the  Measurement  of  H-ion 
Concentration.  l.H  us  uou  return  to  tlie  reaction  or  II  ion  eoneenlratum 
of  snhslanees  in  solution.  As  tin'  slan.lar.l  of  neutrality,  l-ure  water  is 
..|„,sen  I.el  us  eonsicier.  then,  liow  the  laws  of  mass  aetion  ean  he 
,,,,,,11,,,  i„  „,.|,,,  ,.,  ,.„„|.le  us  to  .letermine  the  llion  eoueent ration  ol 
pure  water.  It  has  l.e.^n  state.l  ahove  thai  eliemieally  pure  water  is  in 
,..,,,,,1,1,.  „f  eon.lnetinK  the  eleetrie  eurrent.     This  however  is  not  strietly 

tl„.  e.ise,   '..r  it  eon.lllels  to  a  very  slijlht  .le-ree.      .\. nllli-  to  the  .lis 

.neialion    .v,.oth..sis.   it    must    theref.U'e    he     ivpresente.l     as    e.uitainni- 

mnhr.hs  of  II  <>  an.l  Inns  .,f  II'  an.l  OH',  aii.l  a. r.lin-  to  that  of  mass 

.•i.-lioM  thiiv  must  I...  a  halaii.M'.l  r..aeti.m  helw.'eii  the  inoleeiiles  an.l  ions 
IclU'.'S.'liteil   thus: 


II  ().-=!  II"     Oil',  iir 


lll-l  ■  |oil- 
III."! 


Sine..  \\u'  .•..! niralion   of   II-  an.l  (Ml'   is  ..xtrem.'ly   small,   there   must 

alwavs  he  sueh  an  ovrwh-^lmin-  prep.ui.leraliee  ..f  ll,<>  m.'teeules  that 
„„  ehaii'-'.^s  in  th..  e.m.-eiitration  of  II-  ami  Oil'  will  he  .-apahle  ..f  appre 
eiahly  air.'etinu'  the  .•oii.-eiit ration  .,f  llJt;  in  other  words,  one  may  omit 

th.'   .lenominator   of   th jualion    an.l    write    it     |ll-|   •   lOll'l     -  K.      II 

•■    11  w..  know  the  value  of  K,  it   will  only  1..-  ii.M-.'ssary  to  nwasure  tli.^ 
eon.-eiilration  of  <illnr  W    or  <Mi'  iu  .n-.ler  I..  exi>ivss  in  numeri.-al  terms 

the  r.-aetion  ..f  th.'  s.,luti..n.     It  has  1 n   foun.l  that  the  value  of  K  i- 

ahoiit  1  10",*  aii.i  sill....  th..  eoiieeiitiati..iis  of  II-  and  Oil'  are  li... 
essarilv  e.|ual   in   imr..  wa1..r.  it    folh.ws  that    [III        [Oil]  n  Ix  10  ", 

i  ,.  ,,,i,|,  i„„  |,„s  a  ,.,.n....nli'atioii  ..f  1  l(l^  This  in..aiis  that  water  ....n 
,Mins  approximat..|v  1  -.Mam  m.,1.  .'a.-li  of  II-  ami  Oil'  ions,  .u'  1  firam 
11-  an.l  17  tirams  Oil'  i..iis.  in  U)'  »v  KMHKMMKI  liters.  A  eonseiiueii.-. 
„|-  Ihc.  ahove  law  is  that  no  matt..r  how  iiiueli  lh..  e.uieentrati.Mi  of  ..le 
i,,n  is  inereas...!  hy  a.l.rmu'  another  suhstan.'e.  the  s..luti..M  must  stiP 
,.,,i,iain  some  of  thr  oth..|'  ion.  Thus,  in  aei.l  soluti..iis  ,o,i.  II-  mn-' 
ii„.|.,.;,s,.  Aii.l  .-...  oil'  must  ,|,.<-reas,  in  sii.-h  prop..rtioii  that  the  tu 
,„„l,ipli...l  toiirlh.r  equals  ahout  1  HI".  77m  II  io„  ro„r, , , Initio,,  .-..■ 
h.    us,<l   thir.fnr,    In   i.r,„-<ss   tin    ,;.irtin,i    i,f    ,u„lr„l,   iirl'l   and   olhahr 

sohiliniix. 

Ill    plae..   of   uat..i.    let    Us   -nhstitul..    .h...iiioriiial     hy.lr.x'hl.irie    a.-i 

,„„„l!.  ;,„•;„„,,.   ,1...   >,:U.l    M.-.,..ln,«    ,n    ,„„..    ..f    i.   ...us.    1..     n,„U„.lu,l    l.y    1 '^   .n    .T.U-r    I 

'Iffirrr  "t  'liltition. 


IIYimCMllIN   IDN    CONCI  STUATION 


1(1.1   .V  llCh      llial   is.  11  tiy.lr.M-lil.Mi.'  iK-i.l  solution  .•"iilaininu  on. 


lar  wfitflil    <'l'  liv.liorliloiif  arid   in    «vains  . 


ol'  till'   inolccu 

lit.  !•  ol'  wat.-r.     Al  lliis  illliition 

the   ll-ioH   contM'nt  lilt  ion 


ti'iitli 


lissolvcd    ill   a 


lll'l  is  !M    piT  c'lit  .li-soi-iatcil:  tiuTrfoii' 
Itcn    lor  sliort  i    is  O.K'.H    .V, 


lor  I'll   as  it   IS  wri 


,r.  ill  iiiatlifiiiatical  notation,  11.1      !<• 
Mdhiiil  of   /'.'.//in.ssMif/  (',,.     To  avou 


s('\cijil  tiiiiircs  to  cxpifss  t  I 


(  ,,,  as 


1   tlif  iifffssity   of  liaviiifi   to  use 

lias  1 II  iloiic  aliovc,  Sinviisoii  lins  intro- 

.hi'.Mii  a  s.-iiV.in.-  liv  'wliicl.  only  on,,  li-iiiv  is  vv,\u\vvd.  This  li..'iin-,  <l.'s- 
i„„,,l..l   l.v    l'„.   is    foiiiMl    liy    siilitrartin-    lioni    tin-   pow.T   oi    t.-n    (i..-.. 

ti„-  li.Min-'staii.linti  I.Hiiii.l   Hh   tin-  .•"in n  lo.„'aiitlii '  Hi-'  li'-'iin-  ('x- 

,,,..sMnu'   llu.    marity    ..t'   tlu-    a.-i.l.*      1m    a    .i...-iii..niial    IK'l    soli.t.o.i. 

,l,,r..r..iT.  u.-  must  sul.tra.-t  fnun  tli.'  I.onv.t  -J.  tl...  .•.uuiiioii  lo-  ol  .'1, 
ul,i..l,  is  .-It;  (as.M.|tain.Ml  IVoni  lo..Mritl,ni  tal.l.'s^  Lmviii-  1.<»4.  .  laU.- 
;,„„tii.T  .■Nanipl.-  .l.rinon.ial  ac'ti.-  a.-i.i  is  .lisso.-iat.MJ  ..iily  t..  tlu'  ."x- 
,.,„,   „j-  ,  ;.  ,„.,,  ...,„t  ;  (■„  is  tln.nfor,.  (>.(>t)l;i  ..onnal.  ..r   l.:i     10  •-     M'"''' 

th..  L.-aiitliiii  ol'  l.:i  is  .11,  1' 'i"l>^  '■'      H.  "''    -'^"■^  . 

Tl,..  ..nlv  ol.i...-tinn  to  til.,  us,,  of  th..  ..xpon.nt  l'„  as  an  ..x|..vssion  ot 
tl„.  n.i,.ii'con....ntiation  is  that  it  iii.T,.as..s  in  nia-nitn.l..  as  th.-  a.-idity 
|,..,.,„n.-s  h.ss:  this  is  lu..-ans,>  tin-  n.--ativ.-  sioi,  of  th,.  pow-.r  is  .lisy."- 
(rar.l...!  As  Stat..,]  al.ov..,  it  is  usual  t..  .'spiTss  th.'  stivn-th  ot  alkali.'s 
,is  w,.ll  as  a.'i.ls  in  t.Tins  of  C,,.  or  1',,.  h-.-'aus..  it  is  ..asi.T  to  iiioasniv  tlu- 

,.,„i.M.„tia1i r  II   i.M.s  than  of  Oil   ions.     A  IM    XaOH  solution   is  S4 

p..,.  ,.,.„(  .risso,.iat,..l;  th,.n.foir  th..  'M.ll-  ion  is  I..0S4  A'  ( i.  .'.,  0.0S4  -ran, 
,.,|uival..Hts  Oil  p,.r  lit..ri,  ami  sin.-,'  th."  lu-o.lu.-t  of  th,.  II-  aii.l  Oil' 
,..,n.-,.iitrations  must  alway.  ,..|iial  1(»  ""  uit  L'O  ('.',  it  is  ,-l.-ar  that  as 
tl„.  II  ion  iiuMvasis  in  .•oncntiation,  th..  <»11  i-ui  must  iT,-ipio,.all.y  <!,'- 
.•iviis...  K\pi<.ss,.,l  iKM-onlinj:  to  llu-  ahov..  s,-li..m...  th,.  <).()S4  A  NaOII 
solution   !:iv..s  l>„    1:!.1H;:  thus,  ().(tS4       S.4      Kt--.  tlu'  h.-.   ..f  S.4   is  .!»'J4. 

an,1  this  sulitiaft...l   from  th,'  i>o\\,.r     'J        l.HS  as  I',,,,,  of   14.14      l.OS -= 

lli.dfi  as  I*,,.** 

Siiiiihiriy.   I'l,  of  ("l    A   Ml,ll<»  snlution   is    ll.-JSC.      Its  ,liss..,-iation   is 

1.4  per  ,'<.nt;  tht.iTror.'  th,.  s.iliitioii  itains  only  ().1»I14  i-iam  ...luivalcnts 

llO-i..'.,     1.4     10       r„n-=:J    0.141;      •2.sr.4     .-.     l'„     14.14    •J.s.-.4  = 

ll.i'Sd.t 

•Stri>lly   N"«kn.f.    I'l.    i-   Ih.    \..^:n«h:u   ..,   t!„    I.:,,-.     !M  ..f   tlu iitr,,ti..n    .f    II    i..t,-   in    t;r.„„. 

t't-r    lilrt.    til.     ruk-ilivt     -n;n    tuini;    iiiii|<  r^t."i,|, 

»ll  «.  »i-h  t..  ixrnv,  tl„.  v:,l„>-  .,f  I'll  ill  ordinary  ii..|.iti..-i.  «i-  inu-l  liti^l  111.  .i.itil.iKarMl.m 
..I    ilM    .lilT.  r.ii.-.    li(t«..Ti   ,li.-   v.iliH-   "f    I'll    nii.l   111.-   ii.M    liiflur   wlml.'   luniili.T;   <■.  >■  .  ;l..l"        '■*■;■ 

111.-   anlil..uanthMi    ..f   ".55   l.i-i'iK    '55.   Ih,-   I'l.    i-.    .V55  ■  M  ".    .-i    ". .ii"il,n,15.>    .\  ,   ..r    .-.-■'--    Km.    »>■''■ 

II  i.iti  in  luti.nim.iH'n  lii,  i*. 

■•It  nniM  li.-  r.-iii.-ml..r.-.i  tli  .t  111.  |...«.  r  ..(  •.  iiunil...  i.-.li.  .i'.  -  tin-  iiii.-.!..  i  ,,f  tir.i.  -  Iv  «lmli 
llMl  Mmiil..r  ni„«t  1..  im,Mi|.li.  ■!  I.v  l.ii;  tli.i-.  I'm  '  ,!.„«  .,..1  nu.ii  'I'-"  .1'":,  ' '  "'•, '\''W,!',l'i -.  " 
than  |.|i-'.  I.nl    1   v   HI  \   In  x    In  x   In  x   in  x  In,  ,,r  I  .nnil.llilll  tiru-s  i.  s-..      Sn-;ilarly,   I  il  »  i^  I"""  tnii,  - 

.-IS  KT.al  a-.   I'll  ".  ii..t  nn..-  a-.  Kr.-at.  ,        ,        ,    .■         •    ,    i  i      i      ,  ,.  ;,     ..|-". >  I:.,--. 

A   .,,:iili,.n ..n.niK  .,lii..  -I   .,...:lk    I    ,:r.-,;-..  Tr.-;Iic-,:ii-  ^'f   iii>--.  ri..-l  !.,  .!..■>;•  ■■  n-    .n ..-.. 

■^H,*.tiliit. -.  a  ii.-iitral   •-<Jiiti"ti    i  I'li    -  71. 

JTlu-  .x|ir.~si..i-.-.  I'll  .111.1  I'll  may  l.i-  nsi-.l  iinli-.criniinati-l>-.  Inil  wlitii  tia-  luimiriial  yalui  is 
Xiy.-n.  It  is  n.nst  r.mv.nutil  In  iiso  the   fornuT. 


•_N  riiVMiiHiii  MK  \l,   li\M>  HI-'   ni\,s|iiliM;irM,   iiiiii  1  ssis 

Application  of  the  Law  of  Mass  Action   in   Determining   the   Real 
Strength  of  Acids  or  Alkalies.     W-'  Iniv.-  m'.m  ili.ii  it  is  lli<-  .l.t-'r.c  ..f 

.lis>(priilli..li  ll|...ii  uhirh  tl'.r  Ifiil  si  I. 'lis.'!  Ii  of  all  aci.l  .|.'l..'li.U  iiIkI  that 
this    X.iri.'S    V\illl    .lilllllnll     ,|M-.'     \'.t<.       'I'l Illilil'lilllll     ln'twr-ii     \\m'    uii- 

.lissocial.-.l  iiii.l  iliNN.M-ial.'.l  iii.iIim'uI..,  iiiav   tli.T.'fi.ic  I..-  stiiri.'d  in  lilli.'r 

,lii-.cti..M   l.v  .-lianjiinu'  Hi iniil  lal  i<>ii  ;  in  ollnf  \\..i<ls,  tin-  pn-.Tss  <.l' 

(lisMiciatioii  is  a  i-.'\ci>ilpif  r.M.'tii.ii.  ami  may  In-  ifpi  csciit.'.l  as 
AT.  tt.\'  I!-.  'I'lir  lau  111'  mass  actimi  milsl  aiiph  in  siirli  a  r:\^r.  ami 
IIS  a  niatliT  ni'  i"a<'t   it   liiis  Im'i'II   fmiml  tliat  a  .■mistant  can  Ik-  cali-nlatcd, 

wliicli   is  kmmii  as  llir  .li-.s,..-iali.ii iislanl.*      ll   is  an  .spn  s>ioii  of  llic 

inli.'tvnt  al'ililv  of  ili.'  a.-i.l  to  .lisso.-iat.'  into  ions,  ami  is  llifivfor.'  IIk- 
iii'^l  iiM'asur.'  of  llic  sln'n.:lli  of  lli.'  a.-i.l.     Tliis  is  stii.'lly  tlif  rase  -for  all 

of   llii-   wcaliir  ariils.   Init    sti ;:   liiiiii'ial    aciils    (ami    liases")    do   not    ;.'ivc 

a  salisfarloi-y  loiistanl.  so  llial  llic  roinpariMin  iiiiisl  not  In'  liiaili-  lii'twi'i'ii 
lliciii  ami  wrakiT  oiios.  Tli.it  llir  ilissoi-iali •onstaiit  expresses  the  rela- 
tive slreiiL'tli     of  oru'anie  aeiils  .'an  lie  shown  liy  eoiiiparinv'  its  value  with 

lli.it   ,,|    ihr   lal.-  at    uhieh  e: su^jar  is  inwrteil    i  M'e  pai;e  "J:!*,  tliis  lieinu 

propoilioiial  to  the  eomeiilratioii  of  the  II  ions  present.  K  for  some  or- 
■_Mliie  aei.ls  is:  Aeelie.  ll.dlKldl N  ;  Konnie.  0.(10(1211:  lien/.oi.-,  O.OOOOtl;  Sal- 
ieylie.  (KIOKrj. 

•■|-h.-  ..|„,.liM„   „     ^    ■'■  k.    «!..,,     ,.    1-   .M|..".'i   ll:..l    I"    >"lni..'    1     ■■(   "'      -I'l 1    I'lT,     is 

1    tr,,Tn,,rr>;,I,  1,1    ,'l    !  I.  .  II  Mm.  .     .n.l    li,,l    lli.     .I.l-.,,     ..1    .11 i,.i...ii    I-    .1      H.     .|i,-|ii'it>    i.f    n.i.li. 

>..,  1,,1..|    ,1,  .1I..IU.     .|;il,.|    V    .1    !•(. t    a    tiMin u  ,1.  nt     mil    I'f     1    1.    .•■"■1    tin     .in.llililv     lI    .  .1.  ll 

I. .11   ,1.      T..   ,lln-lr..i..    i.l    11-   l-.l-''     .ll'^    .I'l.!    Ill    >.iii"'i-   .lilnti'.ii-: 

V                                             a  K      1"^ 

,,..,.,4                                             ii.iiH  I  1..' 

.  ,,.  1.1  I  1  ^-^ 

,; .,'  ii.ii.,i,  1 71. 

is.i  iM'irs  ii.rt; 


(  IIAI'TKIJ   V 


1 


» 


riiK  njiNcii'i.i'.s  iwoi.vKi)  i\  tin:  MKAsruKMr.NT  ok  tin: 

llYDUtMlKN  Ht\   ((iSt  r,\TI{.\TlO\   iCmit'.li 
THE  METHODS  OF  MEASUREMENT 

The  Electric  Method 

III  onl.T  to  iitKlfistiiiMl  iIk'  piiii.-iplc  .it'  tlic  >tiiii«l;ii.l  111.1I111.I  \isi'.l  I'c.r 

iinnsiiiiiii;  tin-  II  ioii  fipniTiitriiliipii.  it   is  11 ssarv    tliiil   ;i   l<u    wnitis  l>i- 

siuil  i-otii-fniiii'.'  the  r;ictiii>  i;<'\ ci iiiin,'  the  development  of  electric  cur- 
rents in  cliciiiii-iil  l)jitt<'ii(s.     TliciT  iiiiiy  lir  si  tiiitliir  iippiiriiliuii  »\'  this 

kiiiiul.'il>.'c   ill    1- -tioM    witli   till'   u.'iicijiti.ph    of   tlic   I'liTtiii-   .•iiiniils 

«iii,l iMiis  iluiiii','  |.liysii)lc>'_'ii-  ii<'li\  ity,  iis  in  nctivc  'jIiiimIs  iiikI  ninsclis. 

Whni  ;i  iiiitiii  is  imiiicrscd  in  a  s.iliilinn  <>r  mii'  i>\'  its  sails,  it  lias  a 
tfii.lfiir.v  til  iri\r  n(T  ii.iis  iiitii  thr  si.liitii.n.  Siniilav  ions  aic,  iiow  fX  cr. 
.iIiiskIv  prcM'iil  in  this  solution,  an. I  lli.sc,  l..\  lli.ir  ONiiiotii'  prcssiiic. 
ti'iid  to  oppose  llic  passai.'!'  of  thi-  ions  coiiiinir  I'roiii  tlic  iiii'tal.  'IMn- 
loicc  «ith  uliii'ii  till'  niflal  s.'iuis  out  its  ions  into  tin-  soliiliou  is  i-alh'il 
Ihi  ilirlnihilir  siiIiiUdii  jnissiin.  If  this  is  riplal  to  thi'  osniolii-  pirs 
suii'  of  till'  iin'lalli.'  ions  in  tic  solution,  thrrc  \*  ill  he  no  cjii-liii-  riiiii'iit 

Ufiiriali'il.  hut   if  it   is  iiii'attT  or  less  than  the  os tii-   ini'ssiirr  of  tin' 

iiiiKiilif  ion,  an  cli'i-t  I'ic  curifiit  will  lir  set  up.  NVhfii  tin-  solution  pres- 
suiT  is  till'  nii'atiT,  till'  iiii'tal  will  hci-oiiic  iii'uat  ivrl,\  chaiiii'il.  hi'i-aiisc  its 
ions  (Miiy  posili\c  I'hari.'is  into  tlu'  solution  ii-ationsi;  on  the  roiiliaiy. 
ulirli  the  osiiiotii-  picssuic  is  ^rrraliT  than  llir  solution  pri'ssun',  thr  iiii'lal 
will  have  a  positive  rliarjii'.  ouiii'.'  to  tlii'  iri'ript  of  tho  posili\c  rations 
fi'oiii  till'  solution. 

I'.r"ailsi'  of  a  foiiT  railed  1  h  1  Iriishilii  iillnii-li"ii.  the  ions  triveii  o|V 
fioiii  the  metal  ran  not  travel  an.\'  iiieasiirahle  distaiiee  from  the  opposilel\ 
ehaiiied  mass  of  metal,  so  that  fioiu  one  of  the  eleeti'odes  alone  it  is 
iiiipossilile  fill'  lis  to  lead  olV  any  cleelrie  enirelit.  I'm-  this  purpose  we 
must  form  a  eircnit.  This  is  done  in  the  manner  shown  in  I'iir.  7  h.v 
eoiinvetilii,'  side  tulies  eominj;  from  the  elei-Iiode  vessels  with  an  inter 
mediate  vessel  eolitailiiliij  a  solution  of  liiifli  eoiiduetiv  ity  and  I'.v  eoii 
nci'tiliir  the  nirtals  li.v  wires.  If  the  eircnit  is  fornird  he!  ■,-.  r.en  the 
same  metals  in  solutions  of  the  same  eoiieeiitration,  no  eleetrie  eiir 
lent     will    lie    u'eueiated,    lieeause    the    two    elietrode     potentials    will     hi' 


ni) 


iiiv^n  '•( 


111  \1lrvl     lusis  i.K    illVSlnliMiHM     riliicKHHKM 


il  niiil  ill  ii|)l>ii-ilr  .^H^c■|jull^  tci  .ill 


h  ..till  r.    <  »ii  tlir  .itlifr  liaml.  hIi""!'! 


till'   I'liiii't 


iilnili a    till'   iiirliillii'    I'Mi    III 


III.'   >i.liitiiiiiN   111-   iiiii'ijiliil.   Ill'' 


tl'iillM'tiv  !■    I'liirr    "  I 


I    Itiiu     111. Ill    I 


I, I.,  ilii'limlt'   t«>  111"'  i'IIh'I-.   "II'I 

,1,..   ,„...s„ii.'  uilli   uliii'l,   it   ll.iu^  "ill   I I"^'l   •"  ""•  -lilV."""".'   i"  <•"•' 

..|.,itr,'itii.ii  ..I-  III.'  I""  >..li.li'-t-      Tliis  i>  111.'  |.|-i.i.'i|.l.-  ■.!•  a  n„„.  nlnilxw 

.,11,  iinil  if  ur  kiH.u   tl 1 nin.li.m  of  "• f  Hh'  soliilioiis  ,M.,n|...siii!r 

i,     „„.|    tllill    |.r.i.'r.'.l    t.i M„r   tlir   I'li'.t  n.llH.I  iv.'    lor.-i'.    XV .'   .-1111    .il.faiM 

lli, ,i....|,tniti.iiiH  i.f  til.'  on.ir  si.i.ili.-n   h    .linVn-ii.'.'.     Ti.  .!..  Uus   ^u■ 

nnist  I'lni.li.x    a   loiiiuila   xvlii.'li   tiil<.>  int Il^i.l.^a1l"ll   ll..'  nlati..ii  I"' 

IVM'i'h   till'   iLiti'iitial   aii.l  III.'  .•i.ii.'i-ntiati.-n   ..f  lln'  M.lilli.'ii. 

Til.'  i.i.ti'iitial  i.f  an  uiiUiinun  .lii'iio'l .iiil'nM'ii  of  a  ni.'tal  in  .-'.ii 

inH  xvitli  a  s..liiti.ii.  ..I' ..Ml'  I.f  its  sail,  may  alsn  I ti'i'iiiinnl  l.v  miikinu' 

j,  „,„,  ,,„|,,  „,•  a  liaitiiy  of  u  iu.li  lln'  hIIhi'  i-.l.'  is  i'..iiii...s.'.l  ..I'  a  slaiiii 
anl  I'li'i'timii'  nf  U'-liali'-'inir  Un""!'  l-l-ntial.     An  .l.'.-lnnl.'  ..f  III.'  latl.T 


..,..h..l.      --1     ' ^     -•-■"•'      'V'     -"'"";■■     ,       ""      "''"ll-     "1.'  I        ,      ,     ,n,l,.      ....1..!^ 


.1.1.1. iiii:    11,1..    .lilt. mil     llui.!-: 


Ii-H.iilil.     ....  I.I      .li;  I  111^    ml..    .1.11.  II  III    lli.i'l- 


tx|...  .-an  must  iva-lilv  I..'  iiia.li'  l.>  l..iMi:iMU'  pmi'  nii'i-.-m-y  in  .-.nta.'i 
uith  a  satin  nti'.l  s..luti..ii  ..f  i'al..iii.-l  (ll'-',n,i   in  in-inial  |...tassiimi  .'lil.. 

ri.lc  s.,liiti..n      In.i.r  siiilal.l.'  > lili'.ns  (i..'..  w  luii   tlic  i-in-iiit   is  .'..m 

,,l,t.'.|i.  a  i...t.iilial  ..f  li.r.iiii  V.  is  il.M'li.l.i'.l  in  this  so-ralU'il  <;,lni„.l 
,l,rtn,il<'  tlial  is.  pi.sitivi'  i.ins  „f  in.  iviiry  ai'.'  .I.'|...sit.'.l  .>ii  lln'  iiii'iviir> 
fn.m  th..  raliiMii'l  s..luli.m  al  tliis  ,„i'ssiin'.  Siii.|...s.'  that  « .•  .•i.nn.'.'l  a 
,.,.,l..'n.'l    .•li'i'tn..!.',   Ihriintili    ih.'   int.'iiii.'.liatii.n    ..f  s..ni.'   s..liiti..n    whi.-l 


•  I'll.  ..il,,.ii,  1  ,N,i....l.  .  .1.-1  1>  .  f 
...rl.,|  l.v  .l,..ili-  ...  ..  l.i  ilii.  Ill'  V  .1.  » 
..irr.i.i..ii»  ,1.1. ....!.    Ill  .ii.ii.K.I   M  1  -.In 


:it.ll.h     -li:.;..  .1    vl;.--    \  .«-.  I    ■  "1.1.11. li.iK    |...r..  ^.ii.-r.  ll.  v.    •• 


l.v.l   111.,!    il   :il-  .   nil-   .1   -i'l'-    li.l"-   """" 


t1         .1         ...lkll^.,.lll.    nil..     l.l-M.      I.I'MIIIllI.'.l'-'-'"'*''         

^i-H ';;■■:'•  ;.T:.t.;;;.::v':;'-.,': '-Ill' 'in,,,  1. ,....;: i....^..  i":;;,;^:,,::^:;',,;:;:,-- 


«hi,l.    a,...i..l..iiv    III.     |.r.,.lii,  l.,,i.    ,.f    111,     .1...I"    .ii.ri.il 


1IVI>H<M1I  \   inS    rtiNI  rSTflATIiiV 


:n 


ill  stTVf  iiH  ii   ;.'"' 


tuMJos    Ix'iim    iiN"    •••I 


i|     c.h  .  Mi'ir.     «illl     iiM'illi.T     ilrrlnMlf,    III.'     !"'•     <l 

ith    clfitririil    HpiMiiiilus    fill' 


iiiifflril    Itv     wilt"'    « 


iiK'iiHiii'ini; 


till'  toliil   iMili'Mliiil  III'  lln'  IpiiIIi  i\  ;  ill- 


i.liliim      ti.ttlO  v. 


till 


,„  .„■  Mil.lnt.'liiii:  lliis  vahi.'  Ir.-m  thf  f.tal   i-tr.  r|,.'n.hnu'  on  tii 

MUM    oi    II..'   UllUll.."!!    .■I.'.'II-.mI.'.    XV.-   .'.11.    I.ll    III.'    IH.I.'Mli.ll    .if    111.'    llllkll..WM 

1  {.•.•tliwlf. 
g              W.'  liav.'  .lis.'iiss.'.i  lli.'s.'  i.iiii.'ipl.'H  .il'  .■l.'.-ln..'li.'i.iiNtiy   l....',iiis.'  lli.'V 
S         inriii  111.'  I.nsis  III whi.-h  .l.|..-ii.K  Hf  >n<ii.liir.l  iii.iImi.I  I-m'  the  deter- 
mination of  the  H-ion  concentration  of  fluids.     Sii|.ih.s...   i.,i   .'Siimpl.', 

Iliiit   ih  pin.'.'  ..r  iiMMU  II   iiK'tiil   i"   "" iiMni.-li..ii  ..f  "II.-  .'I.'.'lio.l.'.  \N«' 

UM'  an  .'l.'.'li.Hl.'  .•..n>.i>li.i..'  ..f  a  la.v.T  of  j-uf.'  li.v.|r....'.'ii  k>\-*  in  .-..nla.-t 
I  uilli  a  s..luHo,i  in  vvlii.'li  ar.'  Iiv  II  i"Ms;  tli.n  Hi.'  rat.'  at   ulii.'li  11  i-'ns 


^- 


□ 


*©* 


,,l     III,      H  1..1.     ."11. all"!!  III.'     .'■" 

,  ,  iini:     N,     ■.  I     uilli     Kl   I     -■'il 


1  1^'       «         I'l.ifi.m.     ■■!       irl-ILilM-     l"l      111.       111.  .1-111.  n 
,,,,     t..i.i.h.|     II'     tl.'      I..III.H      ii..>.,l     ..I     ..ll.'lM.I     ,l..ll..l.. 

ii'.i  ",i.i  ,1!.  11  ,i! ;;,  ..I  .h.„  r...i I....-.1 .1 •  - ", "•'  •; "'',■','":,;■',;:= 

i.ii.iu,    „„.    ,,...1,    .11  ,,.,iiMiii.,i ...|.i;i.,..   .I..II 1.1    I-   1—1   '■■  'I'"':.  ",".   ','  *    ... 

.  Mir.  I.I       ,,l      >,.|..,i-      |..,Mli,,ii,      .,(      111.       iil..\.li.l.       ..1111.1.1       ..11      III.       '"I'll- 
-•,,1,  II-,  1,    1 


iM,,.liiH.I      li.ini 


lnMi.nir  aililnl  to  ihi'  M.liiticiii  Ironi  Hi.'  II    lay.r,  nr  lakni   rn'in   it.  will  .1.' 
pi'iiil  .111  111..  . 'Hill-. ntiiiti. Ill  111'  II   inns  in  Miliitioii.     In  oi.li'r  I"  M'l-nr.'  a 

liyilrnu'.'ii  .'l.'.'lii"!.'  fiiKillinu   III.'  alinv.-   i'.'i|iliicMH'nls,   it    is  n ssmy   1" 

.'iiiploy  sDMit'  111. ■.Ills  liy  wliicli  ji  l,i\cr  .if  li\  .linu'i'ii  may  Itc  fiiriiisli.'.l,  ami 
f.iilimatfiy  lliis  i-Mii  I..'  iliiiii.  hy  lakiiii;  ail\  aiitai;.'  of  tin'  pnipiTly  wlii.'li 
Np.iniry  plaliiiiiiii  pnssrssrs  iif  alisiiiliiii'^  lai'^f  .|iiaiititi.'s  .>!    litis  i;as.      II 

is  alsii  I cssary  In  U.-rp  an  alimisplii'i'.'  .'f  pur.'   II   i"  .'inilm't   willi   ''"' 

liiii.l. 

As  is  111.'  .'ii-i'  ..f  Ihv  siiMjilvr  rrils  .ir-crih.'fl  al'"\'',  tlii'i-.'  ai'.>  1\v'> 
•  yp.'s  wlii.'li  \M'  iiiiirlit  use  f.n-  iiicasiirin.:  Ilu-  .'Ic.'lr.'imilivc  f.ncc  ir.'H- 
.'I'ali'.l  in  111.'  unkniiwn  .'l.'clniil.':  a  (      ,'('nlrali<>n  i-.'M  .•nniims.'.l  .if  Iw" 


;V2  rin  sK  (H  NIMH  \i,  hams  hi-'  I'livsinifMiUAi,  cunn-ssiis 

liv.lluui'll  rlcctrndrs,  nl'  xvlliili  niic  cnlltMi  lis  M  si(hlt'ni||  ol'  hliiiini  H  iiill 
i-ciMci'iili',itiiin,  ;iiiil  tlir  cilliiT  till'  Miliiliciii  in  u  liicli  tliis  is  iiiikiKiu  n : 
iiiiil  i\  crll  III'  wliii'li  line  I'li'i-liiiilr  is  ;i  sl;iiiil:iril  i-iiinliicl  clri-l  rnilc  ;iliil 
llif  ulli.T.  ,\   livill-'iiii'ii   flci-truili'  roiilniliilii:  'ln'  iliiioKiw  ii   snhitimi. 

'I'lic  I'Xiirt  jin-inurliiiiil  i>r  till'  ;i|i|i.ir.itlls  in  uliii'li  tlic  i-mIiiiiii'I  rlcr- 
timl.'  is  iixil  will  1"'  si'i'ii  ill  till'  nri-i>iii|i;in>  iiiir  sUi'tcli.  Tin-  liviliofrcn 
I'lci-tiiiili'.  it  will  111'  iiiitii'iil,  is  ii  M'l-y  siiiall  \'  sli.-ipi'il  tiilii',  in  wiiirli  is 
siisp.'ii.liil  :i  |>lnliiiiiMi  will'  I'lmli'il  will)  s|iiiim>  phitiiiiiin  iiinl  ilippiliu' 
ilitii  ,'1  suliiliiili  wliirli  lirarlv  (ills  lilt'  tiltii'.  Till'  spiii-r  illmM'  till'  si)hltii>ll 
is  tillcil  witli  piiii'  li\  ilrnLTi'ii.  'I'liis  ami  tlii'  I'aluiiu'l  rlci'tiiuli'  ari>  I'l'ii- 
iii'i'tril  with  siiitalilr  clri'liii'  iiirasiiiiiiir  iiistiiiiiii'iits,  aiiil  till'  ciiTiiil  is 
I'liiiipli'iril  liv  I'Diiiiri'tiii'j:  till'  t\\ii  rli'i't  ii'ili's  li.\  iMi'aiis  ii|'  all  iiili'riiit'iliati' 
vessel   I'liiitainiiii;  a  Natiiialeil  snliitinii  ul'  pntassiimi  I'lilmiiii'.     Tiiis  ciiii 

lii'etiii','  siiliitimi   i>  iisi'il   lieeailsi'  it   lias  lieeii    I'liiiiiij   tlial   tt leetric  elir 

i'iii1>  -.ct  lip  lit  till'  I'l'iitai'l  lielui'i'ii  ililTei-eiit  snlutiiiiis  ale  sii  small  lliat 
tlii'V     eall    lie    irisli"j:aii|ei|.* 

As    niitlilieil    aliiive,    tile    li  \  1 1 1'litreli    eleelriiile    is    tlial     w  lliell    is    llsi'il    In 

iletei'iiiiiii'  till'    II  i -niiei'iii  latiiiii  lit'  liliiml,  tile   pai'tieiilai'   point    aliiiiit 

it.  ill  eniiipai  isnii  wiili  till'  apparatus  iiseil  I'm-  simpler  snlilt  imis.  Iii'lii!; 
tlial  till'  IimIiii'^i'II  iv  imt  eliaiiueil  in  the  cnuise  nf  tile  e\  peiimeiit .  Tlli^ 
pleeant  ion  is  to  prexelit  tlie  eailiiill  iliosiile  nl'  llie  liliioil  I'lom  lieill'J 
'■  \*  .i^lii'il  iiiit"  ol'  it  li\  a  t'reiilleiitly  elialiiiini;  at  nios|ilii'ie  of  liviii-oireii 
.Maii\  iiiai'i'inaeii'^  in  tin'  earlier  resiills  olitaineil  li.\  tliis  metlniil  were 
due  til  til'  leiiioxal  111'  eailion  ilio\ii|e.  w  lliell,  as  we  sliall  see  later,  is 
one  III'  till'  iliiet'  ai'iils  emit  lilmtiiiL'  to  llie   II  imi  eoiieeiit  ration  of  Mooil 


The  Indicator  Method 

.\v    poii!ii,|    mil    in   a    pievimis   eliapter    i  pa'^e  "JJ  i .   the    iiietliihl    ol'   tilra 
tioii    t'oi'  ai'iililv   or  alkalinil.v    in   wliieli   a   staiiilaril   soliit'oii   of  alkali   m' 

ai'iil   i~  aililiil   until  a  eertain  elianue  in   ll ilor  of  a  siiitalile  iinlieatof 

iv  iletreleil.  i|iie^  lint  affniil  aiix  ill  foinia  I  ioii  leya  II 1  i  ll','  till'  11  ion  eon 
eeiilratioii  ai-tiiall>  pie^enl  in  tlie  solution.  It  lells  lis  tlie  lulu}  fin 
II  III rnliiiii  ol'  avaiialile  aeiil  or  base,  Imtli  ili-^soeiatiil  ami  nmlissoeiateil 
r.\    iiioiliiieat  ioii  of  till'  meilinil  of  pii  I'ei  1 11  le,  liowe\er,  we  may  also  iis'' 

imlieatms    I'm-   iletei  iiiinini:    the    Him iieent  ration.      The    priiieiple    ol 

this  iiii'thnil  ilepi'iiils  nil  the  lael  that  there  are  eertain  ilyes  W  liief 
.•han'je  i|iiite  ill  >l  iiiell>  ill  tint  willi  very  slinhi  i'hanu:es  in  the  llinn 
enlieelit  ratinii.  -Ml  that  if  \i  e  U^e  ilyes  wliieh  piisse>s  this  pl'iiperty  .'.t  .'! 
piiilit    near  that    "i  neiitralitv    i  i.  e..  Iietweeu    rnll..")  ami    I'nSi,   we  can   es 


•\      1. -,  ripli..!.     ..1     ll..      I..l,.>i        l..r     .11    .l-M'li;     I'l.      .I..1.I.      i...l.iili.,l     .l.\.l..],,|     l.\     ll.,-    ,.11     «..|p 

I..      ..Ill     ..1     |l.i.        ll...  S:ll...     I'.    -.IV     lliil    ll..     •.li.ntlli     ..I     111.-    ;iniinl     1 |.|I..|     Willi     lll.il     .1 

....r.i.l    ..1    I.TI..UII    ........ll.    lK..,i-l...|    l.v    :.    1........ ll i,.|,iri...n    1 i:    in,. I.-    l.v      ,    I.m.Ik.     «   . 

V.    ,   ,   .|.ill.irv   .1..  I..  1...  M    11    1..  1.  ..   .  ..pi..*..!    I..  .1,1.    1    1 1..    .1 .in. I   .1.  t 1    .' I. 


IlYPRfKIKN-lOV    <()N(  T.NTUATKIN' 


3.1 


■5 

i 


tiniiitc  tlio  H-ioii  (M.iici'iitration  of  flic  body  lluiils  witli  very  iriiiaikaMf 
acfurafv,  provided  certain  ])iecamioiis  are  taken  to  circuiiivent  tlio 
dist\irl)iii!.'  iii(liieiice  wliicli  tlie  i)ro1ein  and  salts  in  tliese  tluids  may  liavc 
on  tlie  color  elian<re. 

To  iinderstainl  this  nse  of  indicators,  it  is  inii)or1ant  to  bear  in  mind 
lliat  one  solution  reaclin<r  neutral  to  one  indicator  may  liave  a  ll-ion 
concentration  wliicli  dilTers  very  markedly  from  thai  of  another  solu- 
tion reactins;  neutral  to  anotlu'i-  indicator.  This  is  liecause  indicators 
react  to  dilferent  Il-ion  concentrations.  A  solution  that  is  ne\itral  to 
phcnolphthalein  has  a  P„  (.f  alxuit  !t,  whereas  oiu'  neutral  to  methyl  or- 
anjre  lias  a  P,i  of  ahout  4.  This  can  he  very  clearly  shown  hy  titratiiii: 
a  solution  of  i)hosphoric  acid  with  deciiiormal  alkali.  After  a  certain 
amount  of  alkali  has  heen  added  it  will  he  noticed  that  methyl  oranv'C 
chaiifies  from  red  to  yellow,  hut  after  it  has  chan'_'ed  and  is  therefore 
alkaline  as  .iudi-'ed  liy  this  indicator,  it  still  reimiins  distinctly  acid  to- 
wards iihenolphthalcin  (shows  no  red  tint  i  even  tlioULdi  conside.Mlily 
more  alkali  is  aihlcd.  The  melliyl  oranjre  is.  therefore,  itself  unresi)on- 
si\e  to  weak  acids  such  as  remain  after  the  ureater  part  of  the  phos- 
pliorie  acid  has  heen  nelltl'ali/.ed  \>y  '.he  alkali. 

The  series  i.  ;dicalois  which  has  heen  eiuphixed  for  this  purpose  is 
..'iveii  in  the  accompanyinjr  tahlc,  ahui-;  with  the  I',,  limits  throuvdi  wliicli 
they  elianH:e  in  color. 

r.iHT  ity  lM)ic\r<ii:s 


riOMIcAI.   NAMK 

riiv 1    siill'i.ii    plilliiilfin 

(Mil. I   riMiu''') 
'r.ii:i    liniiii"    |ilii'llipl    ^"1 

:   h  plitliiili'lii 

( »rtl iil.i.w    ])«'n/cnc 

:i/.i  .'.,  iiiilii>l 

aiiiliiH' 
Orthii  riiil'iiw    l.t'i)/t>iic 

11/11  ill  I'l.'i'vl 

,'oiiliiic 
iU  liroinn  orllin  cri'sol 

Kiiirmi   |i|itliiilriii 


l>i  l.rniiiii  tliyinnl  snifnii 

|>litli:ili'lii 
I'lii'iiiil  siilfiin   )ilitli:ilcin 
nittn.    rrt'siil    slllfon 

plitliMli'iii 
Tliyinnl    xnlfcm    |i)illir>li'iii 

Orlliii   crcwil    |>hthnlrin 


COMMON 
\  \  M  K 

Tliynitil  liluc 

Itlnni   I'lll'lllil 

liinc 


Mi'tlnl  icI 


|'iii|i\l  ri'il 

r.nnji  r-rrsnl 
IHirplc 

r.iiiiii  tlnini.l 

l.liic 
(■III  Mill  nil 

Cn'siil  ml 
'I'liyiiKil  liliir 

Crcwil 

Iitit(i:tlvill 


CMNCI  S 

COI  OK 

TKATIdV 

rn  \  M.I'. 

/.,,     ,,„l 

e.iil 

K.-.l  yilli.w 

II. Ill 

V.llim  l.lnc 

II. 11^ 

K...1  mIIi.u 

11. ni; 

U...I  yrM..w 

II  III 

Y.ll"« 

|.iir|.le 

mil 

\-,.||..H  Mil. 

(i.o:; 

Vrll.iH  ir.l 

o.o-j 

V..||.i\\  rc.l 

not 

Vrlliin  ri'il 

KAMJK 
I'll 


1.J2.S 

:'..(i  i.i; 

1.1  (1.0 
l.sfi.f 

.-..'J  r..x 


r'nl.:,l. 


fill 

T.li 

C.H 

SI 

7. 'J 

•".S 

S.(l 

'.l.'i 

e  n 

0  >i 

Thr«ir    (Iv««;   ni.iy    fmw   hv    til»tninr'l    in    thi-   cniint'V- 


(W.   M    (Mark  :iml   U.   A.   \ai\'-. 


:u 


iMvsii  (MiirMHM.  iiAsis  iiK  I'livsun  (m;I('M,  ruiMi.ssiis 


I'.ii.th  s1;iti(l  llic  imi'IIkmI  for  ini-Msiiriii','  tin-  II  imi  <'iiiic('iitrati(in  coll- 
-i-t^  ill  |if.'i),niiii;  .-1  series  <>(  sdliitions  eont.-iiniii!.'  known  (•(ineciit rations 
ni   11  ion     lii.il  is  lo  s;i>  .  of  known  I'n     and  addinij  to  caeli  solution  an 

,.,|iKi|    ; unt     of    ;in     indieMtor    w  liieli    exlnliits    easily    distiii<:uislial>l(' 

tdian-jes  in  tini  at  II  ion  eoneeni  rations  aiiproxiinatinj.'  tliose  lielicved 
to  lie  preM'iil  in  tlie  unknown  sohilion.  Tlie  same  indi>-ator  is  added  to 
tlie  nnkno\vn  snhilion,  wliiidi  is  then  |)laeed  side  liy  side  with  tin-  stand- 
ards to  (ind  with  wliieli  id'  llieni  il  nmst  (dosely  niatidn-s.  The  series 
of  solutions  of  known  II  ion  eoneeni  ration  is  prejjared  liy  niixin<r  lit- 
teenth  niOMial  solutions  of  N.i.llIM),  and  Klld'O,  in  varying  propor- 
tions as  ■ii\en   in   the   follow  iu'^   table: 


ri>'  !■  \H  M  !■    N 


I  Wli^HIl     S'.l  I    TlitVS 


ML    til.     pr^'i.f  i..H»   ill. II., 111. I    I.. !..«    1.1   ..I. I. nil    111.-   .I.'..ii..l    I'm: 


■I         ,17 
|i        '.1 


;  ,1     ,M     :.=■     7'.     7. 
■\      1 1      1/-  >    1.1  ■    1 1 


II       S  .'       S  4 
I.       .1..'       .'.II 


r.i      rr      >i      si.'   ..-•  s   si      "i  .;   'u.i  "(..s  "sn 

I  I  I. .11,    i.i>>.    N..\niti<.-  ,111,1    M.ittii.tt.l 

■Sl.ei.'.Li'l    i,li,.-,,li  .1,     iiiiM;n-.~    ,ii .     ;,ii|..i,.l    .i. .     i.Imii;    t.,    s.i.ii-.ii'-    .l.r.-.l; .1-    l.,ll.'«s: 

I     1-     |,    ,1      .1    |.|     ,.i     |.rrii.iT.     |.,,l:i  — nun     |,h.  .-i-li.il.        'I'lTs    ci.rn-    "I     ill.      I'Hl'      I .  .  r> -l.llli/.  .1     «,ill 

iKIiril.,    .,!,     ,li--.,:\.,|    HI    II. -hU    .h.lill..l    «:il.i     .111,1    iii:i.l.     iij.    t..    1    lil.l. 

|"|;,„,.l      .Ik.ibM    ,.,-,  ,..H  I. m    -■,.1111111  I. li.-l  li.it.  111.    l-iir,'   i..rvo  ,]li„,l  -;,ll   I  Xa.lil'n.l  'l|.,(ll 

I-  ,^l..,.,  ,|  I..  ;lu  ail  |..r  (ii,iii  Un  .l:iv<  1..  !«..  «.-.k-,  |.r..ti.l.-.l  fn.in  .Iiivt.  Trii  ni.il.nil.  -  of  watiT 
.,  ,,,,,,11.    ,1 ,..    tn.ii  ,,n    in.l    ,  -,.l;  .,t  111.    I '.I,»  \,i  il|-ii,..il  II  1-  ■■111 1;   II. s"'-  tii.nii- 

1,1    till-  -.1.    ,li-.,h<.l   ill   Ir.-lilv   ili-till..l   »:itrr  .in.l  iii;i.li-   iir  In   1    liti T.      Til.'   soliitic.ri   .^lioul.l   (five  .1 

,l..-,,   r,-.     r.,1   ...l.it    mill   iili.  M..lrlilliali  in.      Il    niilv    a   faint   |.iiil.   i..'..r   is   nl.lairif.l,    llic   sail    is    m.t 

*\!rti.  1. iitK  i-'ii.'. 

Tlie  indie. itor  untliod  is  e\treliiely  ai-enrate  wlien  used  with  pure 
siiliilii.iis  of  .leids.  liut,  as  iiieiitioned  alio\e,  it  is  apt  to  lie  inaeeiirate,  at 
least  willi  iiiosi  iiidieators.  u  hen  protein  oi-  inoriranie  salts  are  pres- 
iiil  ill  the  Miliitioii.  and  ol  eoiiise  it  is  ipiite  uiiilsalile  with  eolored 
llnids  siieli  as  Mood.  In  order  to  o\  eri-onie  these  dinieiilties,  the 
ihdhisis  ,ii<llii,<l  has  reeiiilly  liiiii  evolved.  It  consists  in  iilaein-;  the 
lliiid  jiloiid.  for  e\aliiple  in  a  di.il.vsel'  sae  eoniposed  of  eelloidin  and 
alioilt    as  lai'je  as  a   small   list   tuhe.     The  sae   is   plaeed   in   a   wider  test 

till f   hard    'jlass   eontaininu   an    isotonie   solution    of   sodiimi   ehloride 

llial  has  lie.n  earefullx  listed  to  aseerlain  tli.it  it  is  strietly  neutral. 
Till'  .'1111011111  111'  Mood  or  seiiiiM  i'i'i|uired  for  this  nielhod  is  only  2  or 
:;  e  .-,,  ;iii<|  ihe  amount  of  s.ilt  solution  |)laeed  outside  the  sae  should  he 
.iliuiii  ihr  siiiiie  It  lakes  onl\  from  li\i'  to  ten  minutes  for  dialysis  to 
oeeiir.  The  eelloidin  sae  is  then  removed,  a  few  drops  of  the  indie.'itor 
ail'  thoroiiulilx    miNid   with   the  dial>sa1e,  nnd   the   tuhe  eonipared   with 

the  si'iiis  of  sliindards   until    lln rrespondinii   tint    is   matehed.     This 

indieates  the   ll-ioii  eoneentiatioii   in   the  dialvsate.     The  lints  pioiju I 

li\    iisiiiLr  sulplioiiephi'iiolphthalein  .'ire  reprodii 1  as  nearly  as  possilile 


.1 

IS 

m   i: 


Ph7-i 


P„  7./  P,.  /-a  Ph7-3  h.  /■'<■ 


n 


n 


r      J 


nn 


w 


w 


Ky 


10 


Pn/i 


Pn't 


P„7  7  P.r& 


\J 


\J 


Ky 


\ 

f 

1 

i 

<i 


i 


IIVIi||(((:i;\   llt\    (  ()N(  I  NTIIATKlV 


M 


ill  fill'  ii('('oiii|iiiii\  iiiji  I'lijirt.  Tlic  ll-ion  cinn'ciili-iiliiin  i>(  tlic  iiiiUnciw  ii 
Miliitiiiii  is  tliiit  III'  tlic  lint  with  wliicii  it  iiiatciics  in  tlic  sci'ics. 

It  niiu'lif  lie  tlii)ii<.'lit  tlijit  this  iiicth(Ml  udiilij  he  iiuiciMHiitc  hccnusc  (if 
the  hiss  (if  cjirhoM  <lii>xiilc  Inmi  the  liloiid.  liy  actiiiil  c\|ieiiiiicnt,  how- 
ever, it  has  Iierii  fiiiiiKl  that,  il'  the  lihind  is  cnllecteil  with  ecrtain  i)1'p- 
eailtions,  the  ciripr  is  iicu'liirilile.  The  iiii'thoil  is,  theicrmc,  a  iimst  iisct'iil 
line  elinieally. 

The  fdlhiwiiit;  talile  irives  the  liV(liii<;i'ii-i(in  riineent  ration  or  true 
Ifaetioli  of  the  hoih    lliliils. 


Kl.l'lll 


4 


l:l I 

I    linr 

ii;i-'t  rir   jiiii-c  I  ;i<lul:  > 
liastric  Jiiiri'   ( iiil;oit  ) 
r:iiH'H':iti<*   Jnii-c    (Jhl:1 
SiiimII   iiiti'stiiiiil  rniiii'iK- 
SmimM    ititi-^tili:il   ntnlr'it^ 
r.ili'   fi'nin   liver 
I'.il.    tiiMM  ;;:ill  l.hi.Mer 
!•'  isj.iiallnn 
1*1  I  ^)>)i;it  ioit 
i't  .u  ** 


J'll 

Ki.nn 

I'll 

7.4 

.Muscle  jlllee    i  I'msIi  ) 

(i.s 

fi.ll 

MiiM-li'  juice   1  uiit    l^weci ) 

V:iri:il.le 

<*'.!' 

r:OMTe::s    e\1r:ii'I 

5.« 

II. 'I  1  i> 

l'<'iit<>lje;>l    ll'ii.l 

7.4 

."i.ll 

I'er  iiMl'li.'lt    lluhl 

7.4 

s..'! 

.\ljUei>ll!4     llllMliir 

7.1 

s.;; 

\'itie"Ms   liiiinur 

7.0 

Inrniit  1       -M 

(■eriliri.^|.ili;il   lliii'l 

7.2 

7.S 

('enlii.is|.iii:il  lliiiil 

s.-j 

.",;;  7.1 

.\nntii>li(-  tlni'l 

7.1 

7.1 

Aiiiiiiulii'  lliihl 

R.I 

i.-> 

.Milk   1  liiiiiiiiii  1 

7.11-7.; 

7.2 

.Milk  (ciiw  1 

(1.1!  (i.s 

Mi'  :   (  L'l'iit  ) 

fi.(i 

Milk   iiiKsi 

7.fi 

(\V.   M.  Chiik  :iri,l   II.  .\. 

l.lll.S.) 

I 


ClIAITKIJ   VI 

TIIK    I.MKil  I.ATItiN    <»l'    NKI  TI.'AMTY    l\    TIIK    ANIMAL    r.(»l)Y 

AM)  ACIDOSIS 


Niptliiii!;  is  iiiiiic  cniisijiiit  in  llu'  animal  rcDnoiiiy  than  tlic  Il-ion  con 
niitiatiun  i('i,  III'  Ilic  llnicjs  wliicli  liatlic  the  tissues.  This  rct;ulatiiin 
is  InnciaMicnlallv  ol'  a  |)h\  siccii'lii'iiiifai  nature,  dependini;  on  the  inter 
aelKin  nf  alkalies  with  aeiiis,  ul'  uiii<'li  eailiniiie  and  phnsphorie  acids 
and  the  proteins  are  tlie  must  important.*  When  dilTeient  annnuits  of 
aeids  or  alkalies  are  added  to  water,  the  lan^'e  of  vjiriatioii  in  II  ion  is 
\iM'\  extehsixe.  \\iieie;is  in  Mood  the  ran^f  is  \eiy  limited  indeed,  not 
exteiirlim.'  lieyond  I'nT  ;ind  I'h7..')2  (i.e.,  ( ',,  lU'V  f!'  iro(>s  nliovc  tlint  of  a 
(i.ouii.itiiii.l  .V  sdliitioii  or  lielow  that  of  a  (I  (Hiii.ddil.o:!  .V  solution).  In 
other  \\  ords  IiI'mmJ  c.in  w  it  hsl.-inij  ecmsideralile  additions  of  ;ieid  or  al 
kali   witlmut    mueji  ehaiiiie. 

Buffer  Substances.     The  eliemieal   react ii^ns  upon   wlueh  this  remark 
alile   eniistaiii-.\     in    tiaeliiin    dipends    ha\e    lieen    e\|)l;iilied    hy    Law  I'euee 
.1     llendcrsdM  ''      The    Innijamentiij    i'(piiitions  ;ire   ;is    rollows: 

M  lll'o.     II  \       Mil  I'o,     M  \,  .■iii.l 
MlU'd      II  \        IMO      M  \. 

uln  M   M         :i   l..'i~i  '   i.'i.lirl.-.  .-in. I    A.  :iii  .'i.  i.l   r:elirl<'. 

Now  it  li.is  lieen  ilisco\ered  that  weak  aeids,  like  ('arlionie  and  ])hos 
pliorie,  possess  the  lemarkalile  property  of  maintainin!;  the  reaction 
constant  wiien  they  ai-e  present  in  ;i  solution  whicli  also  contains  an 
excess  ot'  their  salts,  rnder  these  <'ireumstani'es  the  concentration  of 
ioni/i'd  hydnitj-cn  is  almost  e\ai-tl.\  eipial  to  the  product  of  the  dissiM'i.i 
tinii  cnnstaiit*  of  tlie  acid  (see  pa'je  \'.))  multiplied  hy  the  ratio  In 
tweeri    free  .-leiil   aiiil  salt;   in   other  wm-ds. 


II-        K 


II  \  I 


|l'.\|- 
If  eailionic  acid  is  present   in  a  solution  of  liiearlionates  so  that    ther' 

•.\i  .  ..i.liiiL'    !■.    1  ir.  ■  ■n-l.iiii.-.    iiMhn:-    1  i.n    .1.1    .ilhir    .i-    ,,.  i.U    ..r    .i^    .Tik.'lii^.       'I'luv    .irt-    III.- 
I. .11-    ..ill.  .1    aii>|>li..U  M. 

Mil.      .■■IM/  il!..TI    -l.-ii'l    I    .-    .ilii  ;i.!\     I..  .  II    :.  (.  ;i.  .1     I,.    :is    .1    t.LMII.-    ulii.  Il    i  \!ii  ,-...■«    llu-    1.  ii.l.  .. 

.il    .1   wt-.ik    :i(t.|  i.r   i.;i-.-   I..   .ii--<..  L.l<-      in   an   .iipii  ..lis  si.liili.iii.      "h    rv|.r.-ss,s   (he   iir..|..rtl.'ll    in   wd' 

III-     ""111 I.1I..1    |.:iil    I-   ..i|m1.i.    ..(  .M-I.nu    111    111.     |.i,--,iui'   ..f    11-    i.iiiv."   an. I   lli>i.(,.i.     i-.   a   i;.ir 

..f    111.      -I'.Mi.lli        Th.     .1 i.ili,.ii    ,,11-1111    .1111. .11111,    I.,    .il...iit    II  11I1I1.IMI11.5    f..l    r.iil...iil,-    .-1.1.1;    Ih.i!    ; 

III.    .Il--  .1  1  ,11.  I    ..1    11.111;    i.;i..    II     ijllil:'    .,1    r....iii    iiini.i  laliii.-    »ill    li.-    mi.  Ii    that    lli.-    i..ii..iii 

II..II    ..I     II    i.>ll    l.]ll.l!-    .1    iMMiii.liiHi.;     .\    s,.llll|,,|l 

lit! 


ff 


A(IIM)SIS 


37 


.irc  iMjiiivfilt'iif  <|ii;iiititi(s  III" /(•<(    II .« '( >,  iiiiil  liifiiilMiii.-ilr     ir..    ||,4|         , 

till-  ll-iiiii  ciiriri'iiti-atiiin  will  In-  cxiictly  tin'  siiinr  as  ihv  ilissmMali'iti 
■•(instant  nl'  carlMinic  ariil:  llicivfnir  (l.tMHI.tKHI.")  N  (I',,  ti.:!h,  or  aliniii 
livr  tinii's  till'  vahif  of  nnitialit\ ,  (i.(l(i(l,(l<l<l,l  N  (  I'n  7.:>li.  If  ten 
t'niit's  as  iinirli  fn-f  ••arlionic  aciil  as  liiiMi-lionati-  is  pi-rsmt,  tlicn  the  li-ioii 

IMA 


I'linccntratioM    will    In-    lifly    times    that    "f    niMilrality,    i.  c 


1(1 

lUAl  ^   1 

(l.(IOO,0(Mt,r)        ().(>( )(),(»();■)   (I'll        ').:(!);   if  tlicic   is  ten   tunes  less  eallmnif 
ai'iil  than   hicailionate,  tlie  ll-i(Ui  cnMeentiatiiin   will   he  mn'  half  that   nf 

neiitralitv.  i.  e.,        '^  (l.(l( )(»,(« HI..")       (l.(l(i(l.(i(l(l,(l.'>  i  ( I',,        7.".1)  ;  nr 

if  twenty  times  less,  one  foiirlh  (I'm  7.(1'.  Sinee  a  larj-'e  amount  of 
liiearhonat(~  is  actually  present  in  hlooil  (enou<.'h  to  yield  from  ")()  to  fi.'i  e.e. 
t'O.  per  1(1(1  e.e.  of  hliNnh  (see  pav'e  :illl),  anil  the  flee  earhohie  aeiil 
un(ler<roes  llnetuations  wliieli  are  only  trivial  when  emnpared  with  those 
wliieh  have  lieen  eliosen  in  the  ahove  esaniples,  it   is  elear  that  there  must 

lie  very  little  ehantre  in  the  1!  ion  eoi ntration  of  th,'  hi I  in  eomparison 

with  the  variations  whieh  would  oeenr  were  no  liiearhonate  present. 

Aiiothor  weak  aeid  which  ads  like  carhonic  in  maintaininir  neutral 
it\  is  iiciil  plHisi)hate  iMIIJ'0,i,  and  for  the  same  reason  namely,  that 
ils  dissociation   cnnstant    is   of  similar   mauniitude   to   the    ll-ion   coneen- 

ti'ation.     Althou(;h  the  hi I  plasma  itself  contains  niueh  less  phosphate 

than  liiearhonate,  the  tissues  contain  a  considerahle  ninoimt,  which  en- 
allies  tliciri  to  maintain  their  neutrality.  This  action  of  liiearlionates  and 
phosphates  is  styled  the  hulTer  action,  meaninir  that  it  serves  to  (!ami» 
down  the  efTeet  on  the  ll-ion  concentration  which  additions  of  acids  or 
alUalics  would  otherwise  liave.  As  pointed  out  hy  I'ayliss,  however,  a 
hetter  word  to  Use  would  he  "laiii|ion  action."  since  flic  siihstanees 
actually  soak  uj)  much  of  the  added  II-  or  Oil'  ions.  It  is  not  oonlined 
to  the  fluids  of  tlie  higher  animals,  hut  is  very  widely  distributed 
throui;hiiut  luiture;  for  exaniple,  in  the  ocean  and  in  the  lluids  of  mai'ine 
orfianisnis  and  animalcules  (see  L.  ,1.   Henderson)." 

Altliouirh  the  actual  rea<'tion  hy  which  neutrality  is  maintain<>d  is 
purely  of  a  physienehemical  nature,  some  iirovisioii  must  oliviously  he 
iii;ide  so  that  the  acid  and  basic  substances  that  take  ]>ai't  in  it  may  be 
Mipplieil  and  those  prodin-ed   hy   the   reactions   rcino\('d   as  occasion   I'e- 

cpiires.     The  sourri  of  siipithi  is  partly  exofr us  and  partly  eiidojreiious. 

The  oxojionous  source  is  the  basic  and  aeiii  sid>stanees  present  in  the 
f<iiid:  and  althou<rh  we  do  not  ordinarily  attempt  to  control  the  amounts 
lit'  these  substances  iiiirested,  we  may  do  so.  as,  for  exainple.  by  the 
persistent  ndministration  of  so(b\  in  cases  of  pathologic  aci(h)sis.  The 
endogenous  source  depends  on  the  constant   production   in   metabolism 


:{8 


I'livshiM'iiiMirvr.  ii\Ms  (IF  iinsiKi.iMiifAi.  ruoci  ssi;s 


of    iiciiN    slli'll    lis    c'.ii  liiiiiii',    |>li>is|)li()ric,    l.irtic,    niid    Mil|ilimic,    iiiiit    <if 

nik.'ilii's  Mii-li  lis  iiiMiiiiiiii:).     Aiiipliiilcrii'  Milist;ii >..  like  iiiriiim  ai'iils  ninl 

|)i'ritciMs,  iiiMv  fiiiii'tiiin;iti'  titlicr  iis  iii-i<ls  111-  jis  iill^i'iliiw.  ^VIl;lt(•\l'r■  iii.iy 
lie  its  siiiii'fo,  n  cnnsiiliM-iiMr  rrsrr\i>  (if  iilUiili  is  iinilimlitcdly  availalilc 
ill  till'  iiniiiial  cuiraiii^iii.  Tliis  rciiniicil  st'Pic  ol'  alkali  appears  to  lie 
aiilniiiatii-allv   lilinatfil  as  a   result   ul'  tlie  pli vsicuclu'iiiical   pruci'ss. 

Till  n  niiirnl  is  afl'crtril  li\  tlilcc  pal  liw  a.\  s:  ll^  tlirciili.'li  tlic  iiilitrs 
irasi'iiiis  i'arliiini<'  ac-id  is  cliiiiipatrij -.  i  "2  >  tlii'diiu'li  tlii'  kiilncys,  tlic  tixi'd 
acids;  and  ' 'i     tliimiirli  tlic  iiitcsl  iiics.  senile  ci|'  tlic  plnisplioric  acid. 

t'arlHiiiic  acid   is  prcujuccd   in  lai-Lrc  aiiiniiiits  in   ilic  iioriiial   pi ess  <it' 

iiictalMilisiii,  and  is  c\crctcc|  in  a  LMscmis  cmiditiiin  liy  tlic  Iiiiiirs.     Varia 
tiiiii   ill    its  cM'ictiiiii   is   tlic   must    iinpni'tant    iiiccliaiiisni    fur   cnnlnilliiiu' 
ti  iiiitnrnrii  rliiiin/is  in  (',,.      In  mdcr  tu  make  this  clear,  it  may  lie  well  In 
revert   fur  a  mcuiieiit  tu  tin'  pliysieiicliemical  c(|uatiiin  liy  wliicli  carlMinic 
acid    is   cnaMecj    to    niainlain    nciitralitv,      Tliis    ma\    lie    written:      ('i, 
II»(» 

Sal  no 

free  carliiiiiic  acid  tn  the  lilnnd  i  as  liy  caiisini;  an  animal  to  respire  some 
of  ilie  j;as  1.  ur  li\  the  adiiit  ion  of  some  other  acid  (e.  ■_'..  oxyhiitx  rii',  as  in 
dialii'tcsi  which  will  deconipose  some  of  the  NalJCO,  and  l>rodiice 
ll,('<>.      The    increase    which    these    chaniics    would    eniise    in    ( ',,    of   the 

I'l 1  is  prevented  lp\    the  reiiiaikalile  seiisiti\it\    of  the  respiratory  cen 

ler  to   chaimes   in    (',,.      An    incieasi'    which    is    much    less   than    can    lie 

measured   li\    pliysii hcmical   means  stimulates  the  center,   caiisiiiir   in 

creased  pulmonary  \  iiilil.itioii,  so  that  the  carlionie  acid  is  inimeijialclx 
elimiiiMtecl  throii'^'li  the  luniis.  'I'his  elimination  does  not  stop  wlicii  tin 
old   lc\e|    of  ciirlioiiic  acid    concentration    is    reaelied.    hut    proceeds  until 

...  II  (•(» 

the  ori'.'inal    ratio    »-,,,,    i>    auain    attaiind    in    the    hi I,    and    < 'n    is 

i-cslored  e\a.tl\  I,,  its  ori'rinal  \alue.  If  it  stopped  at  the  old  CO.  eon 
ceiitration,   the    ratio   would    he   too    lii'.ih    liecailse    tlieie   is    less    N'alK'O 


molecular  rati 


'rile  ratio  ma\   he  increased  eithei'  liv  addiii'. 


THE  THEORY  OF  ACIDOSIS 

Allhoii'ili  these  coiisiileral  ior  ,  indieale  that  variations  may  occiir  in 
till'  hicarlionate  conteht  of  the  lilmid  without  an>  siirniticant  (diaiiu'e  in 
(',..  they  also  show  that  the  hicailionale  coiileiit  must  he  a  I'ritcriiui  of 
the  acid-liase  lialalice  of  the  lilood,  and  plolialily  of  the  hoily  lliiids  iii 
ireinral.  As  |ioiiiti'd  out  liy  \  an  Slyke,'-  liicarliouatc  represents  the  c\ 
ceSs  of  I  se  which  is  left  o\  el'  after  all  the  //'((/  acids  have  liecn  ncii 
!rali/i  d  It  represents  the  liase  tha'  is  a\ailalile  for  the  iieiit rali/atiou  oi 
any  cnccss  of  such  acids  that  iiia>  ajipcar  a  measure  of  the  reserve  ol 
"liiilTer  siilistaiK'c""  or,  more  s| ilically,  tin  nlktilim   rtsirn  of  the  hody 


Iiidir  iinniiiil  iMiiii|itiiiii>  (III'  iiiM'iiiiil  III'  \iilH'(»    ill  IiIimhI  |iI,isiiiii  is  \i'i-.' 


sIjiIII    I  illllnlllltill'_'   til   "I'l  \'>'>    Mils.    \<l-V 


•III  ('(»),  mill  u  Inn  it   is  ii'ilin 


it   iinliratfs  lliiit  iiii  I'Xi'css  oj"  tixcil  .-iciil  imist   lie  picsciil.     'I'liis  is  liiUiii 

li\     Villi    Sivkc    mill    otlllTS    til    ■■nllstitlltl-    till'    r.'ill    lii'lillitiini    i>l'    ./ri'i/i.v/s 


II.IMICIV,        il    •■IHII 


litiiin   ill    wliii'li   tin'  ciiiic'ciiliiil  imi   nl'  liiriiilinii.'itr    in    tin- 

1,1 1   is   ifiiiMMiI   IpcIow    til.'    iiiiiiii;il    liM'l."      ir   till'    i(s|)iriiliin    i-i'iilin 

I'lir  miv    rcjisdii   slnnilil   imt    irspciiid    iniiiipl  ly   (innnjli    tn  iin    iin'i'i  ;isc    in 


tin'  iiiiili'i-illiir  latiip 


I  !,(•(» 
NalH(» 


ami  <',,  ciiiisi'iiiii'nlly   In iiii'  yi>'ati'i-,  tin' 


.•(.mrniiiii  is  i-allcil  II  Ilium  fn  iisiil  III  iiiiilusis.  Inil    if  llir  i'i'iiI.t  ihii's  ris|iiiinl 

sn  tliat  ('m  is  li.'M  I'l.iistant    (  allliniit:li  Nalifd     is  lii'i'ifascil  ..  lli inlitii.n 

is  line  of  I  1)111)11  iixnli  il  iiiiiliixis. 


]■'»]•  iiraftii-al    rcasniis 
I'lnls  till  an  I'sliiiialinii  oT  tlic  Kirailmn 


llirn'ripfi',   till'  sillily   nl'  |iatlnilii'_'ii'  ai-iilnsis  <li' 
ale   (•iint«'nt    nl'   llic   lilnml   nl',   ■.iiii'i- 


il   is  siiiiiili'f  In  carry   nut   ami   is  nl'  niiial   valin',  nf  llm   plasma.      Winn 
iilasiiia  is  nlitaini'il  liv  li'iin>\  iiii:  liinml   I'iniii  a    \i'iii  nl'  tin'  aim  ami  i'«'"i 


liil'li'.'im.'  imiiii'iliati'ly  mil   nl'  i lai-t    willi  air   is,,  i|i;it    (  n     iiia>    imt    Im 

InsI    I'inlil    ill    it    cnlltains  apiiln\illialrlv   t'lO  vols,   lici-  I'i'llt    nl'  CO..      Sit 

II'-' 

;i4>. 


kiinw  thai  till'  partial  pri'ssurc  cif  <'<>.  in  M 1  is  npial  In  (J  mm 


asi'crljiihril    t'rnm    ili'ti  riiiiiialiniis    i 


il'    111,'   alvnlar   (  ( » .  i 


I  M'l'    pa'. 


ui'  can  calculate  Imw   milcli   nf  llic  CO  \ols.   per  cent    must    le   in   simple 
snlnlinii  liy  applicatinii  nl'  llic  law  nl'  snliitiim  nf  uas  ill   a   liipiiil   i  pJi'-"' 

:!:!('.l.      It    lias   In I'miini    lliat    plasma    al    lieily    temperal  lire    ami    at    TlWI 

mm.   Ilir  (almnspli 'lie  pressiirei    ilissi.jves  (l.'i}    per  eini    •'<>..  s,.  llial    at 

4"' 

4-J  mm.  it    uilldissnivc   „  „>  \  !""•  n '•  ■>  I        '■'<  \ 


III) 


tlie  liiriires  t(i  our  eipialimi   «c  irei 


pi'l-    cell 


I.    'rraiiseriliiii',' 


I  I  (  •<  I 


IN'alK'O  I       lilt  '-'It 

Til's  .Icliiiilion  111'  aciiliisis  leave,  mil  of  rc'ranl  all  cninl.tii  n.  liiat    may 


aise  ilic  ran 


ll,C<» 
N'alU  <• 


aiiililinll    nl'    ll.t'O      uitlnillt     ileeomposill'.' 


ali.v   ol'  tlic  N'alK'O,,  sueli,  I'or  csainple.  as  occurs  wlieii   an   e\e( 


il    !ri 


earnoiiie  ai 


ill    is   p 


I'll!    ill    the    liliioij    plasma.      Since    increases    in    Ine 


rt»,  arc  lint   iiirrciiucMl   in  Imlli  lieaitli  ami  ilisiasc     e.  ;:.,  aspli.\  \ial  con 


ilitinlis       llic    all 


live    (IcliM 


llinll     is    lint     sullicielit  ly    compi'ellcllsiv  e.        \\ 


w  e  ennie 


to  .study  tlie  control  nl"  llic  ri  spiratnry  cenler,  we  sliall  see  thai 


an    increase   i 


11    I  he    rat'. 


Il,<'(>; 

NalK'O 


il'    siillicicnl    mai;nilinle    to    cause    an 


actual  increase  ni  (  n  can  he  | 


lie  prndiiccd  hv  causinv  an  animal  In  respire  air 


liltiii.iillv    »itl,    111.     i,-iilt 


.,tt>l    h.Mii    lli>     I,,, 


riii,~.  Il  ».   tak,   1  II 


K    .1-    4.)      Ill  •    I  Ml.  I,.,ili»   ;iii.l    K. 


11. i>,    M,    k:,  1 


ll.l.lli    ■    11  .(>    ■     III 


.V.ilKii, 


4(» 


•|IV>|I  IM   IIIMICAI.    HAMS    (IK    I'llVMllKKilrAr,    I'llOI   I  SSKS 


ii 
ii 


(•iilit;iilliM'_'  nil  r\ri'ss  ul'  (  (  I .  ;i  line  liciiliisis,  luit  Ollf  for  wllifll  no  pllU'f 
is  t'oiinil  ill  tlu'  :il>ii\('  ili'liiiitjnii. 

\c\  iitlicli'sN,  Will  Sivkc's  ilcdiiitidii  luis  ii  iriil  viillH".  liiTiiilsi-  it  ('ill- 
|ili!i>.izcs  tlic  iiii|iiii'tiiiici'  III'  ;i  (Ictniiiiii.itiiiii  "I"  llif  liiciiiliiiiiati'  lis  a  cr' 
li'iiiiii  (iT  ilic  (li'iiici'  III'  III!'  Imiiiis  (if  aciildsis  iiNiiallv  iiict  willi  in  disease. 

Tlic  liicarlHiiiatc   iiinli'i'  Midi  ciUKliiiDiis  may   I miic  rcdiii-i'd  citlicr  lic- 

caiisc  (if  the  a|>|icaialic('  ul'  iiii|ii  (i|i('il.\    (ixidi/.cd   t'alty  acids,  like  /i-iixv- 

Ipiilyric   and   ; tuacclic,    w  lien   carlidlivdiat)'   nu'talMilisiii    is   upset    as   in 

dialii'tts  (If  slat  V  atimi.  (ir  liccailsc  (lie  acids  |ii'iidiicc<l  liy  a  iMiliiiiil 
iiii'laliiilisiii  arc  iiiadc(|iiatciv  cliiiiinatcd  liy  tlic  kidneys,  as  in   nepliritis, 

Acciirdiir.'lN.  if  the  rcs|iiialiii-.\  iiicidiaiiisni  and  inerease(|  mass  iniive- 
iiieiit    ipf  tlic   111 I    (  f(ir  an    increase   in   ('h   aecelcrates  tliis  alsii)    slidiild 

fail  Id  cjiniiiiale  CO.  ((iiickly  ciidiiLfli  sd  as  td  keep  llie  "    ,    '    faliii  at 

diic  lucniictli,  then  (',1  will  rise.  'I'liis  is  not  likely  to  happen  nntil  a 
laiu'c  pail  df  ihc  Nalh'tt  has  liccii  used  up,  sd  that  an  cstiniatidii  (if  that 
aetnall.x  prescMl  must  he  a  icliahle  index  of  the  prdxiniity  t(i  this 
cdiiditidii. 

A  siist.iiiied  iiicicase  in  i'„   is  ii iiipatilile  with  life.     Tlie  N'alM'O.  is 

the  liufl'er.  the  facldr  (if  safcl.\  which  prevents  its  (icciirreiiee.  Alth(iiiu;li 
it    is  did>    in  arterial   lildud    i  i,  e.,  after  eliininatidli   (if  excess  (if  ( '<  >^   l.y 

the  liinirs  has  lieeii   accdiiiplislied  I    that    cdiistaiicv   in   the   ratio  ;,      ' 

Nallt  <>. 
can  he  cxpectcil,  it   is  fdiluiialc,  for  practical  reasons,  tliat  venous  lilood 

ciiileclcil  diiriiijr  ninsciilar  rest  and  uitliout  stasis  slioidd  lu'  only  sli^htl.V 
ditTelelit. 

When  aciiis  are  added  to  the  hlnod.  tliey  will  lirsl  of  all  he  neiitrali/ed 
li>  the  "liiifl'eis"  of  the  plasm.!  iiaiiicly,  NalK'O,  and  pr<it<'ill,  as  we 
li.!\e  seen.  Hut  this  is  only  the  first  line  of  defense  aiiaiiist  lipidosis,  for 
Iniffir  siilistaiice.<  present  in  the  Cdrpusclcs  nia.\  also  he  used.  This  intra- 
ciirpiiscular  reserve  of  alkali    is  iiiohili/.ed    i)artly  h.v   traiisfereiipc   of  K 

and   Na   froi irpuscle  to  plasma,  luit   mainly  hy  that   of  IICI  from  the 

plasma  into  the  corpiiside,  so  releasinir  liase  in  tlio  former  to  eoiidiiiie 
with  the  added  acid  le.  f.'..  ll_.('0  i,  aceordiiifr  to  the  ('(platioii: 
ll.fo  \;i(!  «  \allt(»  IICI  The  IICI  on  enterinir  the  eoriMlsele 
reacts  with  phusphatcs  aecdrdinj;  to  the  e(|iiatii>ii:  IICI  !  \a„HI*0,p^ 
Nail  r<>,  NaCI.  This  is  a  particularly  iinportaiit  detail  of  the  Iniffer 
action  of  the  hlood.  t'or  it  shows  us  how  tlie  plios))liates  of  the  eorpuseles 
are  reiKJered  availaldc  for  nciltrali/iii!,'  iicids  added  to  the  plasma,  wiiere 
there  are  praelicall.\  no  phosphates.  Indeed  tin-  transference  of  acid 
throiiy'h  the  i-i.rpns<u!ar  eti\<!"pe  indicates  that  the  same  Miit  of  lliiii^ 
must  K"  on  with  the  othei'  cells  of  the  liody.  so  that  the  plasma,  itself 
rather  ixior  in  liiitT»r  sulistaiices,  has  all  those  of  the  liody  at  its  disposal. 


.\(ilN)SIS 


41 


THE  MEASUBEMENT  OF  THE  RESERVE  ALKALINITY 


1.  Titration  Methods 

Tlinc  IMC  scMial  iucIIhmIs  hy  wliicli  llic  ns(  rvc  nlkiirmity  nf  I  he  lilooti 
iii.iv  Ik-  iiicjisiircil.  Tile  siiiiplt'st  iti  llifiiry  fniisists  in  sn-iiij;  1m>w  ninrll 
siiinilnnl  iii-id  iriiist  Ik-  jhlilcd  to  a  iiiraHiircil  ((iianlit.\-  of  I)I(km1  plasma  in 
iiiilfr  fii  rcarli  the  nciilial  point  a-  ,iu>l)i<'il  \>\  "lianifc  in  tint  of  soiiii' 
iiiiliralor.  The  in<lir,.iuis  cniploycil  (r.  jj.,  nictli>l  oiani.'i'l  art-  sui-li  as 
rliaiii.'"'  flit'ir  tints  at  J  ion  conccntiations  tliat  aic  wfll  to  tlio  acid  sidf 
lit'  neutrality  (i.e..  at  a  liiirli  <'ii  <'f  low  I',,  i  To  Ininjr  fin-  i)la,siiia  to  this 
point  oi'  nrutrality  tiir  addi'd  aliv.di  will  iici'd  to  nrulrali/c,  not  onl.x  tin* 
liii'ail(onatc  of  tin-  plasma,  l>nl  otiici-  a<'id  liindins;  sulistani-i-s  as  uidl. 
This  will  irivt'  ns  a  falsi-  impression  of  (he  acid  l>indin<_'  powers  of  til*' 
iilasma.  siia-r  '^t  the  normal  ( ',,  of  the  lilood,  proteins  do  not  altsorli  aeids 
to  anUhinjr  lil<t'  tin-  extent  the\  do  at  hiuher  deifi'ees  of  t',,.  Another 
oliJeetioM  to  the  nu'thod  is  that  tin'  proteins  intei  I'er-e  willi  tlie  sensitixc- 
iiess  of  the  indieators. 

The  ohjei-tions  i-an  he  removed  l>.\  determininy;  tin-  end  point  eleetro- 
iin'trii-ally  or  li.\'  indieators  thai  ehantie  tint  at  aliont  I'm?.  The  most 
praetieal  way  is  to  determiiu'  the  idiantre  in  ('n  prodiieed  l>y  addinj;  a 
known  volume  of  stamiard  aeid  to  hlood  plasma.  The  resiiltitiij  ehanire 
ill  t'li  will  then  lie  greater  flie  less  the  alkaliiH'  reserve.  In  the  ideetro- 
Tiietrie  method  iire^ularities  that  miudit  he  eause<l  liy  vaiialile  anniiints 
of  raihonie  aeid  in  the  Mood  to  start  w  ith  are  liest  controlled  liy  removiii}? 
the  CO    from  the  plasma  aftei-  addinir  the  stainlaid  aeiil      The  jirocednre 

therefor insists  in  mixinji  1  e.c.  plasma  with  2  c.i-.  \  ■")()  |i('|  in  a  small 

separatintr  funnel,  \vhi(di  is  then  evacuated  so  as  to  remove  the  CO., 
after  whi(di  the  lluid  is  transferred  to  a  iiydrojien  electrode  and  (\, 
iiiea.siired  (see  pajie  '1'.)) .  In  normal  lilood  this  shoidd  he  10"'"  ( I'li'i.til. 
Ill  acidosis,  where  there  is  a  tlepleted  alkaline  leserve.  the  2  c.e.  of  acid 
will  cause  ii  much  j;reater  ehannc  in  Cn-in  diahetie  Mood  to  lielow  fi 
"f  lower. 

The  tcchnic  involved  in  the  aliovc  method  is,  however,  too  exactint;  for 
routine  clinical  work.  For  smdi  jturixises  the  colorinietrie  method  of  la-vy 
and  Kowiitreo  may  he  em))loyed. 

Tin;  MKTiion  oy  Iii:vv  .wn  I{o\vntui:i:."— A  ti'st  tiil>  made  of  hard 
i'  iionsol'"!  j;lass  of  aliout  '2t)  c.c.  capacity,  contaiiiin<r  ahout  a  irram  of 
|po\\dcred  niiitral  |iotassiiiiii  oxalate,  is  tilled  with  newly  drawn  Mood, 
•  •v!ii--liaf(-!y  stojipcrcd  aiid  placet!  oil  it-e.  Quit^titie^  i-f  '2  c.c.  *-acli  <>f 
I  lie  Mood  are  then  jilaced  in  a  series  of  seven  small  (nonsol  test  tid)es 
and  allowed  to  stand  for  fuo  to  six  minutes  in  order  to  permit  a  iiari-ow 


4-  l'IIVsrrtM||i:MH'\l.   Iivsis   hK    l'll^^ll>t  iNili'Al.    I'tiiMlsi^l.S 

la.M'i'  III'  pinsiiiii  tn  M'piirtilr  on  ihr  Mirliifc  illiis  iiri'vcntH  lakiiiu:  i>(  t\w 
IiIiiihI  (|iiriii|r  the  suliHi'i|ii)Mit  ihlilitioii  lit'  ill-ill  HI'  iilkiilii.  'I'lu-  lilimil  in 
till-  liiNt  tiilx'  is  iiM'il  I'lir  llii'  ili-lri'iiiiiiiitiiiii  nl'  tin-  iii'iniiil  II  inn.  In 
fiH'li  of  III!'  ni'vt  lliiTc  tnlirs  iiri'  iiililfil  icN|MM'iiM'ly  ti,|,  ii;J  unil  (i :{  r.i-. 
\  ."((I  MCI.  iirnl  III  till-  last  tlirn-,  Niiiiilar  i|iiaiiiitii'>.  ,<(  \  .">(l  NaOII.  Al'Irr 
invi'i'tinu'  tin'  fiilx*  so  as  I"  mis  itn-  mntcnN,  )lir  IiIimkI  in  <'ai'li  is  trans 

•'••iTcd  ti Iliiiilin  .sacs  ami  tiir  r,,  iliici  iiiincil  aci-iiiilinu  to  llic  nit'tlinil 

tli'si-rilivil  i'lsi'\«  hi-it>  I  pajri-  .'fjt. 

Tin-  liilii's  a?c  iioffil  ill  «liifli  a  flniiiu'i'  in  lint  I'l'iiiii  lli.il  iif  tin-  nnriual 
hliMiil  is  i'\ii|int,  ami  tin-  n-siilts  ate  vx|mi'ssi'iI  as  llic  re.  nf  N  ."lO  III  I 
or  \a«i||  wliii'li  must  In-  ailili'il  In  lijiiiiil  til  i-lianiii'  its  (*,,.  Thus,  llir 
iill,iill  Inill.r  is  till-  .M-.  (if  N   .'lO  Nadll    whii-li   lan    lif   aililnl   to  'J  ivc    nf 

l>l'H«l   will I  i-lianj.'!'  nf  ('ii  nf  till'  ilialxsatf,  ami  tin-  nriil  liiilhr  Ihi-  c.c. 

nf  \  .Vl   Il<l. 

'I'lic  iiii'ttiiiil  siilTcis   frniii   till'   fiillnw  iiii;  ilraw  liarks: 

I.   \°<'i'.\    small  i|naiitilics  <<(  ariil  ami  alkali  an-  I'liiplnyi'il. 

■J.  It  is  nfliMi  ililliciill  In  ti'll  jiist  cxai'lly  w  lit-n  a  sliu'lil  ililTcii'iin-  in 
linl    has  Ik'i'h   iiimlui'i'il. 

•I.  Kvni  with  llii'  |iri'i-aiilinns  ili-si-iilii'il  almxc.  it  is  iiii|inssilili'  in  lir 
sine  that  Ihr  aiiiniinl  nf  ('(1.  in  tin-  ililTiimt  saiii|ilcs  nf  lilnml  is  tin-  same, 
"hii-li  mi'ans.  nf  rniiisr.  that  sninr  lilniuls  v\  ill,  nti  this  ai'cinint  alnnr.  In- 
alilc  In  liiml   niiin'  alkali   than  ■illirrs. 

'I'lii,  Miriniii  (IK  \' \\  Si.VKi  .     .\  mi'tlinil  liaxd  nii  smiirHliat   the  same 

piiiici|i|i'.  lull   which  is  iiinic  accni-alc  I aiisc  it   niccls  the  alxivc  nlijcc 

linn,  is  that  siiu'ircslcd  U\  \'an  Slykc  Stillman  ami  t'lillcii."  I'lasiiia  is 
frccil  nf  (■(».  Ii\  placiinr  it  in  a  \aciinin,  ami  is  then  mixed  witli  an  e(pial 
viilniae  nf  .\  ."((»  Iin  (nr  .  0||.  iinil  the  ("ii  (letei'inimd  liy  the  eleeliie 
Mielllnil    I  see    paye   ".Ml » .       Ill    llie   ease   nf   iinrmal    lilnnil.   ,if|ei'   Sllcll    an   aihli 

linn  nf  acid,  a  piactically  nnrmal  ('i,  will  he  fniiml.  w  lieccas  in  Ihe  lilnn.l 
nf  cases  nf  acidnsis   it    will    lie   xciy   dislinelly    incleased    i  i.  e..    I',,    Inwer 


2.  CO -combining  Power 

The    aliiiNe    nli.jeclinlis    In    the    lill'atinn    nf    lilnnil     pl:isliia    nr    dialxsal' 

with  slamlaid  snlniinns  nf  acids  .-nc  reiiinxcd  if  we  nieasiire  the  cnin 
liiliinj:  pnwer  nf  tin    hlnnd  alkali  Inwards  carlmnic  acid  itself  at   iiniriial 

hlnnd    rcaelinll.      This   ina.v     he  d itiler   in    lilnnd    illllllediately   ;iftcr    il- 

reimival  fmiii  the  animal  nr  in  lilnnd  that  has  heen  first  nf  all  saturate.l 
nutside  the  lind\  with  cai-linnie  acid  at  a  partial  pressure  ('(pial  In  Ilia! 
exislin<„'  in  the  limly.  Since  for  practical  reasnns  \cinius  hlnnd  must  Im 
used  ill  the  clinic  at  least  the  fnriiier  nf  these  methnils  siitTeis  fmiii 
the  fault  that  varyin^r  amnimts  nf  carluinic  acid  will  he  aihled  tn  lie 
lilnnd    (hirini;    its    jiassajfc    thrnii^h    the    lissnes,    and    Ihe    eirnr    thereh.x 


AIIIHIKIS 


43 


iiiiMiiiril   will   Ik nil'  LMtiills    (luuriiMilnl   it'  SfiiiiiiH  nIiisIh  Ikih  licfli   pro 

iliici'il  in  ih'iiw  iiiu:  tin'  N|)r<'iMirii  I'ur  iiniilv^i>.  I'lut  iIh-  fjiji't'  i-riisun  w  li\ 
tills  niclhiMl  liiis  nut  Imcii  r\tcii>i\rl\  t'in|)l"\  f<l,  as  pniiilcd  iiiit  liy  \'nM 
SK  ki',   is  tin'   ti'clinii'al    ililliriiltv   nl'   liillKiiiif   the    ni'rcssiifv    nllill_\sis. 

It  is  nicist  siilislartiirv   I Ilfi-t  m'Ihmis  IiIihkI  al'tci'  a  pcrinij  inix'  limii' 

at  h'asf  1  ol"  iiiusiMilar  rest  isn  lliat  lliffi'  is  im  cxrfss  nf  CO.  I  aiiil  witlmiit 
vciiiiiis  stasis,  aiiil  t"  <"fiitril'iiu'<'  witlmiit  pci'iuilliiitf  any  consiilcralili'  Inss 
III' oii'liiinic  ai'iii.     Till'  laltcf  pi'icaiitinn   is  nri'cssary   lircausi'  tliiM'f  in  'i 

iiiinratinii  nl"  ai-ii|  railirlrs.  c.  ir.,  IICI,  riuin  plusiiia  iiit irpiisrics  uln-n 

till'  ^'^y.  nt'  IIm-  t'luiiirr  is  inrir  I'll,  ami  in  tin-  icvfisc  iliicrtinn  when  tin- 
( '(».  is  <li  ri'caHitl.     Il'tlift  I    IiIkihI  well-  niit  till'  saini- ijiliini;  ccM- 

liirnirini;  as  it  is  in  tin-  '•  •!       '  if  scpa ulastna  wiiuiil  nut  i-iinlaiii  tin- 

saiiii'   aninnnt    nl'   all\al       i  i       i'  '  ilihity    wnnld    In*    alti-rcij. 

.\lllliiin;ll    lIli'iirrliiMJ'  .■.;'iilr,     ••■i.l '  ' !''  IM  slmnlil    In'    JiiM  jnrilli'il    in 


-h. :  .j-"- 


"A/ 


If     111       lii.iKi.iiii    1. 1    .ii'i' ii.ilii.    I.ir    -aliii.iini^    I'l I    "r    pLi-iiii    willi    .\|im'1     to         11"'    wl.i-- 

'f.iiU    Ml    till-    )...ltt.     i.ii.I,M*<     t  -. i     in..t,tiii>.       'i'hr    ^1  I'.lt.ittltfc:     ttimul.     .i-     -■••n     ,i-     tl     I.  •>     !■»  r  n 

t>;!|.|     with     .'|'11..|     .111.      .Il..ll,|     I.,      ,|,.„.|     l.»      .1     M.-IIH      .ml     111.       -I..| k      IIMM..!      ..It  li        -     111'" 

t.i.il.-l      ...      III.. I      Ih.       Iil.n.rl       l.ilU,,      ,,      l.hll      ,.,,      11-       vi.ilU. 

:iii  aliiiiis|.lii'ii.  i-iiMlainim.'  tlic  saiiic  partial  pri'ssnii-  of  ( '<  t  as  exists  in 
llii'  liiMi\  II.  c.  the  ahnilar  aiii  i  sci'  paL'i'  :>44l.  this  lias  ln'cn  Inniiil  iin 
jitartiralili-  t'lir  iifniTal  use,  am!  is  uiinrfcssarv  il"  loss  nl'  •'(•.  rrmti  tin- 
spi  I'iiiun  111'  liliiiiij  is  picvintiij  liv  allnwiii"^  it  1"  ll"w  into  tlir  s\  linir'' 
^i-v\  sjiiwly  iwilhont  any  sniMioiii,  It  is  inixi'il  in  tin'  syi-ini;i'  with 
|iii\mIi'I'I'iI  inriitiali  potassium  oxalate  leiinntih  tn  make  a  1  pel-  eeiit 
^iiliiiiiiii  with  the  |iloiii|>,  ami  iiiimediatelv  ilelivereil  into  a  eentiiruvre 
iiilie  iiiiili-r  paralliii  oil,  w  hieh  liy  (loatiii'^  on  its  siir  e  siTves  to  ilimiiiisli 
lire  ililTiisioii  of  ('(».  to  the  oiitsiile  air  leveii  tlio.      i  sin-li  oils  dissolve 

re  CO.  than  water  i.     '{"o  mix  the  hlood  with  the  oxalate,  the  syrinu'e 

sliiiiilil  lie  moved  liaekward  and  I'lii-waid  se\eral  times,  Imt  it  must  not  h" 

.luikell. 

.\ner  eeiitritiii.'inir,  alimit   :!  e.e.  of  plasnui  are  remo\ed  and  s.ituraled 
"illi  CO^  at  the  same  lensimi  as  in  alveolar  air  (i.e.,  .'i..'"i  per  cent  i.     Tlii^ 


iii 


-3 


44 


•liYsir(i(lli:MirAI.  iiAsis  OK  |'||VSI(ii.(mj|(AI.  i'ii(iri:ssHX 


is  (Iniif  li.v  iiliiriii','  tlif  pliiMiiii  in  .-1  sc|>aiiiliii;;  funnel  of  :1<I(I  c.c.  ciiparity. 
lavini:  llif  fiMMifl  i>M  its  sidr  ami  ilisplaciny:  tin-  air  in  it  liy  alveolar  air 
scfurcil  !)>  quickly  niakiin.'  as  deep  an  inspiralinn  as  pussililc  llirou^li 
the  lulii'  and  Ix.Illi'  rontainint;  '_'lass  licails  i  Kis;.  Illi.  The  '.'lass  lii-ails 
ri'iniivc  t-MTss  i.r  walci'  vapm-  linni  I  lie  air.  Tlie  tunnel  iniisl  lie  restup- 
pered  liel'iire  tlie  end  i>(  tin-  expiiatinn,  sii  that  no  ontside  air  enters,  it 
is  then  rotated,  tor  ahout  two  Miinutes,  in  s\ieli  a  way  that  the  i)lasnia 
forms  a  film  on  its  walls.  If  it  is  nreessary  to  posti)one  tlie  saturalinK 
of  the  plasma,  this  shoiiM  lie  pipetted  olT  from  the  eorpusides  and  pre- 
served in  hard  irl ass  lest  tidies  eoated  with  parallin.     Kroin  ordinary  jjlass 


li.'     I  I        V  ..n    SKI..  "-   .11  I  .n.ilii-   l..r 
It 


\nii'^    lilt     t  M    ,  ..nil.inMi^-    i'.iVM  I    1.1    111. Mill    It)    li! I    pl.t^ni 


i'Miiii<_di  alkali  is  soon  dissoU  ed  out  to  \  itiate  the  results.     After  saturation 

of  ihc  pJMsiiia  with  ( '<  •  .  the  funnel  i>,  ph I  in  the  npiiiiht   position  and 

the  plasma  allnwecl   to  inJIiiM    in   the  iiiitrow    portion,  after  wliieh    I    <'.e 
is  reiiiu\ed   with  an   aeeiiiale   pipette  and   anal\zed    for  t '<  >^. 
The  analysis  iii,i\    he  done  li\    iisiii<r  rithei'  the  \'an  Slyke  oi'  the  Mai 

ilalie  liarefiifl   appalatlls        tin     \'iiii   Shil.i    iilillniil   is  ils  fiilliiirs: 

Tin  apparatus  is  IIIIimI  to  the  |ii|)  id'  the  j.'raduated  tiihe  with  Miefeiirv 
I  l'ii_'.  Ill  liv  laisiii'j:  the  iHtreiirv  reservoir  F,  eaie  heiiiir  taken  thai 
/>  and  K  are  also  tilled,     thie  e.e.  of  the  ('(>_,  sat ni'ated  plasma  is  then  dc' 


1 


AOIHOSIR 


45 


livorcd  into  .1  (w!  L  li  lias  lic.'ii  viiisc.l  out  with  CO.-froc  aminoniu  water), 
and  the  stop.-o.-k  /  tiirnv.l  so  tliat  liy  cautiously  loworinc  tlic  levfl  of  tin- 
i-fscrvoir  h\  tlit>  plasma  runs  into  /.'  (l>ut  no  traco  of  air^.  Tlie  same 
procedure  is  repeated  wilii  1  e.e.  water,  so  as  to  wash  in  all  <.r  the  plasma, 
and  (iimlly  (»..')  e.e  of  .'i  per  eeiit  II.SO,  is  sueked  in,  after  whieh  stopeoek  / 
is  turned  olT.  The  reservoir  F  is  then  lowered  sul1ieientl>  to  allow  all 
of  the  mereury,  hut  none  of  the  hlood,  to  run  out  of  /.'  an<l  V.  A  vaeuum 
is  thus  i)roduee<l  in  //  and  V. 

As  the  level  of  the  mercury  falls  in  /.'  and  <\  the  plasma  elTervesees  vio- 
lently,* heeause  it  is  exposed  to  a  vacuum.     To  he  certain  that  all  traces  ot 
Co/have   heen   dislod'ced    from   the   solution,   the    apparatus   is    inverted 
several  limes.     To  ascertain  how  much  CO,  has  heen  liherated,  stopeoek  // 
is  now  turned  so  as  to  hrin-:  ('  and  E  into  conuiiunication,  and  hy  eautiou.sly 
lowirin^r  th.-  reservoir  the  Huid   in  ('  is  allowed  to  run   into  the  huUi   E. 
Stopcock  //  is  thereafter  turned  so  as  t(.  conned  ('  and  /'.  and  the  reser- 
voir raised  so  that  the  mercury  runs  into  C  as  far  as  the  fO,  that  has  col- 
lccte.1  in  the  l.uretle  will  pcniiil    it  to  jro.     After  hrinjrin^'  the  level  (d"  the 
mercury  in  F  to  corr.-spond  to  that  in  the  hiirctte.  the  jrraduation  at  which 
this  stands  is  rea«l.      It   irixcs  the  cc  of  Cd,  liherated   from  the  plasma. 
Iiider  the  ahove  eonditinns  normal   plasma  hinds  aliout   7.')  per  cent   of 
its  v.ilume  of  CO.:  therel<u'e,  since  the  total  capacity  of  the  pil)ette  is  '.(> 
cc  ,  the  mercury  should  sliind  at  tt.i'.T")  e.e,  on  the  Inirette.     For  accurate 

iiicasurenn'nt  it  is  n ssary  to  allow  for  the  CO,  that  remains  diss.dved 

iii  the  water,  etc.,  as  well  as  for  l.aronietric  pressure  and  lemi)erature. 
riiis  is  I.est  ilone  l>v  llu'  us.-  of  a  lahle  l.ased  on  the  known  s(dul>ility  of 
CO,  uniler  the  various  conditions  olil.-iinim:.  which  is  niven  in  Van 
.■sjyke's  paper.'- 

Thf  llnhUini  liiirinifi  ,ii>i>iiriitii^  that  is  most  suitahle  for  the  ahove 
,n,,l>sis  is  shoNu.  in  Ki-  V'X>.  l-a-c  :!vj,t  On,'  cc,  ol'  CO,  free  :iiMMicMiia 
u;ilcr  is  idaced  in  the  holtle  and  the  1  (•.<•.  of  plasma  delivered  heneatli  it. 


•n^i-    tii.iv    111-   iilrvtnli.l    1\    .l.Miril!   .1    «ii.;ill    .llii|.   "1    •.ipTvll.     ..Ii"ll'l.       ,         ,    ,. 

.  rius    (.,nn    ,.|    n.il.l.ui.    l'...ri...o    .,i.|-iri.li,«   1-    n.l    .lUil.-    ill.-    ~:im„.    .„    ill.     .I.tl.  ■  .^nti..l    m.in..in.l. 

,i..,|   f ,....„M,.u.,l    ..I    II..    I',...!;.!.* i;    r"».i    ■■'    I...,..kI.;I.m,    M'W    ■'"    '       '"    tli.-    I..ti 

,     lh<-   i.r,s,„l    ,,„rii..«.-    a  -i.l.'   I'll..-   ..t   111.-  l..-n.l   ..I    111.    I    ml..-   i^   ...MM..I..I    Willi   .1   sm.ill   ml 


.i|    I 


l,.,K.    »h..!i   .•:.>,    U ...SM-.I    l.v    a    -i.w,      Wh.n    111.    .■-■.-    -    .-v.-lv.  .1    in    111.-    l."lll.-.    U    rn-rs 

n   llM-  lli.i.l   11,   II,.-   i..MM,„.-..l   l„„l,   ..I    ,1,.-   ,ii,i„..„Ml.r   .-...r.- „.I,mkIv    .1,1.1   ra,s,s   ll,.-ll   ,„   ih,-   .li.M 

.  Sin..-  II,.-  ...I., ,1.111. 11  ..1"  111-  a„l..„„t  ..f  l.•:,^  .v.lv.-.l  .l.prn.K  ..n  lin.ll,-K  ll,.-  /•.■■t.l.r.-  |ii...l,l,-.-.l 
,..,„   .,„v    ,l,...,k..    „.   s.,l„,„..   It   1-    „...s>;„v    .,1,,,    ,1..     .-.,-   I,..-   I...„   .^.■^•-l    I-   ..".pr,"   'I""    "il'l- i; 

uMl.l  111,  ,„.„1M„^  ..f  ll,„.l  ,„  ll,.  |.,..v,iii,.l  l,i»l.  "I  ll"-  „;.,„"„„  I.r  H  l.r.i„>;l,t  Im.  1.  ,■.  iK  ..i,k„,.i1 
I  Til.  h.ithl  ,-it  »l,i.l>  ll,.-  tl,.,.l  -'.„'l-  1"  ""  'I'^'-'l  I""''  ""■"  "l"i""~lv  c..rr.s|...„.l«  I.,  ll,.- 
--If.    .r-,il..l   l.v    ll..-   .v..l\.i|    tM-i.  ,      ,       .,. 

ih,    ,,,„.„ i„,   .l,„,„„„„.^   ll,.    ;i," ,1   "I   v.,-   .V..U..I   .i.,..„-l-   ..„   ll,.    n:,-  l.i».   «li,.      -1  .■.-•■ 

,h.    i-,.,.,,,,-    ,.l    :i    „..s    1.    i„v.r..h    ,„..|...,i,..„.,l    ,..    ll-    v.-l„i".     il-,>::      "■  ^"V\ ll,..l    ll,t 

.     ..f    i::.<    .v.,lv..|    v,.!-    i-.|„nl    I       ll,.-    »..l„,l,r    ..f    It,.-    l,',"!.-.    ll,>„.    m„.  ■■    H,.'    V..I,,,,,.-    I"'    '"■•'' 

-l.-„,l.  Ilir  |.i.-.„t.-  «..„l.l  I,.-  .l..-,M..I     tli.-il    K  Ih-  «»'-l   i""'f  -I'"-''  Imil.  «"„M  •■■|,i-«l  lh-i< 

.,,„.,M'l..i.-.   "r    M.CH,  „„„.    .f    «.,IT  ..r    1.,., I    .l..^.■      ,1.    «l„-li   <•    H"'   «;'"l   ^u  „.ilH    "---I    '" 

N.    ■„a„..n,.|.-r        Ai,v    ..ll,.  r    ..l,..-,  V  ,-1    |.r.— ,r,-    «..„M    ,l,.,.l,.,.-    ,  .-r,.  s|...-i.l    I..    ,1,.-    v..l„,„r    ..I 
■  '.1   pis  a,-.  ..r.linK  t..   tlir   .  .i,,:iti..tt. 

\..\.  ..I   I...I1I. 

|.,., 1,111   I  wli.  11   .li.v.-  ..il   i"    ll-...!) 


,  ;i„.!    I,,l.,,ii.-   !'.    t„.- 


v   ll,'-   .li'l'.,!.,!,,*   it,   ll 


.-ll- 


,1.1.-1    I 


k.-l    ,„    III. 


ni  nil  .r 


„K 


r   l.v    a   M.I, I'll-   iiianipiitali- 


.,f  ll,.-  I.'ill!.  -  i<  rmil-ii..l.  .iinl   ll,.    I.ilario 


4(i 


fllVSU  (MllI'MIl    M,     |!\SI> 


OK    I'llVSKll  IHJ(M      rUlK  I  SSI  S 


If   liiililc   is  tlii'ii   .•iimiccli'il    with    til.'   iii.'ii irl.'i-   wiili    the   inciMiitiiiiiN 


Tl 

ilcsciilifil  (  Inch  lici  .•   in    this   miIiiiiii'.      W  I 

1 11  mII.iu.'.I   Inf.  s.-itiir.-ili'il  t.iit.iiii'  ,'K'iil   is  iiii\i.|   witli  ll 


icii    li'lii|>i'i':itlli'i iHlitiiitis   li:i\i' 

IC     |llilSIM,'|     Slllll 


tiiill  .-lllil  till'  UMs  cmiIm  .1  liii';isiii'(i|  \t\    the  ilispljiiM  iMi'iit  (if  till'  lliiiil  ill  111 
iiijiiioiiift.T.      'I'lif   ;i|i|>jir.iliis    iii;iy   ;ilsi.    Im'    iis.'.I    \\ii|i    MimhI    in    pliii-c    ..I' 

pl.'ISIIM  III    tllis  Cilsi'.    1 


lllHCMT,   it    In  ni'('css;ir\    tll.lt    I 


ir  <i\\  "Ti'ii  he  ffiniiv  I'll 


re  ;i(|iliMLr  tin-  tjirtiiric  jici 
.1 


I      Tlii>  I 


>rii-;iiit  iiiii  is  II 


III  .iisl(Pili:c  s.iiiic  111'  111.-  (I,  ri-niM  li.iii.ii.'l(i|iiii.     Till-  III. III. I  is  tl 


■<i'Ns;iiv,  SHI.'.'   iiciil 


Is  t  Mcicl.irc  lust 


III'    ,'lll    l.'lk.'.l    W  itil    Illllllliilli; 


I  iMiutjiiiiiiij.'  s.iiiic  sii|Miiiiii,  til. '11   sliiikcn   w'.fh 


"■J.")  (•  <•    -iitiiijit.'.l   |>ii|;issiiMii   t'.'iricyiiiii.l.'  s.iliit ion,  ;iiiil   liiiJilly   with   ill.' 
s;itiir;ilc.|  ;i.-i.|  s.iliitinii.      IT  lil.m.l  is  iis.'.l.  th.'  .  >!  imathuis  iiiiist   lie  inaili' 

11(1    strii-llv    licsh    III I.    siiii'c    .m    stiiiiilinu:    th.'    (  o Iiiuimr    p.nv.M' 

'_'lC!ltl>    ({••ti'iiii|-;itrs. 

3.  Indirect  Methods 

Th.M.'  ,ii-.'  s.'\t'i;il  .illit'i-  iii.'lli.i.ls  li\    uhi.-h   till-  iillv.'ilili.i   i'i>.'i\i'  m,       In- 
lllrilsllll'.l.       Tlloi'    ill. -III. Ic: 

I    Determination  of  the  Tension  of  CO    in  Alveolar  Air     im-;.   .■!44i 

^" '''1^    iiii'lli.MJ    is   I'lniilnM,!    111. ire    |.;ii  I  i.Mil.iily    in    iiu.'sti'.Mliii!.'    tlif 

liiii-iii.iii.'   .■iiiitii.l   i.r  till'    II  siMi-^it.ii'y   I'i'iiti'i',   \\f   sh.-ill   .1.  r.r  .1   ,|.-M'i'i|ili.iii 

"'    i'    'iii'il    l;itiT.      'I'll.'   iilv.'Miiir  ( '(  •     t.'llsi.ii ir.'sp..ii.k  t.i   Ihc   ('(I.   ten 

■-I.III  ill  iiiti'ii.il  liliiiiil  ,111.1  this  is  pr.ipiii')i.iii;il  til  th.'  .•ilk.iliiic   r.-,.'i'\ ,.  ji^ 
il.'t.Miiiiii.'.l  li\    W'lii  Sl\  i..''v  III.  th. Ill  .iN  is  pi',,M'i|  li\    thi'  I'^i.'t  lliMl   Ihf  r:itii>, 

pl;iNiii;i  ( '(  I 

I         ,        ,  ., ,     .         .       ,  Is  >.;it|sl:ict.iiih    .•iihNl.'ilit. 
:i\\  I'lil.-ir  (  « >     ti'iisi.iii 

•J    The  Measurement  of  the  Acid  Excretion  by  the  Kidney.     As  miirht 

I..'  i\p.'cti'.I,  thi'  ;i.-iil  li.is.'  I'ljuililiiiiiin  III'  the  Imily  m.-iv   ;ils<.  !..>  iriumn!  Iiv 

»>,.isin;  nil  III    <.<:   tin    mi. I   •nnlmii    ,,i   th,     iirlii, .    in    whi.-li    tli.'   in-ids    -.tv 

.-'.iitiiiiiiil  piii-tl\    i iiiliiiMtiuii  u!lh  .■iiiiiiiiiiii;i  .11-  ii  lixc.j  lijis.',  ;iiii|  p;iftl> 

III    M    t'lvc   st.-iii'       \Vi'    nIi.'iII    iii'si    ,,i    .•ill    >ii|.'r   th.'    111. ■Ill, Ills   iif   .'i.-i,! 

.■\.'ii-liiiii    iiii.j    Ih.'ii    .'N.-iiiiiii,'    till'    .'V  i.l.'ii.'..    shiiw  ill.,'    that    th."    tntal    a. Mil 

«'\.-i-.'li.iii    is    pi-.ipnilii.ii;il    til    ih.'    .•ilk.'ilin,'    ri'scrvc    .ms    iiwaxiii'.-.j    hv    th.' 

.•llii.v  «•  .li'N.-tilii'.j   tiiith.iiK 

Km  1(1    THIS     111-    .\.   11.    |\     C.MI'.IV  Xlluv     Willi     AmMiiNM.       'I'll.'     p|-(nlllcti.ill 

"I       ailllH. .111.-1       IX      .■xM-lllla|l\        ,,,;       .'ll.j.iir.'ll.lllS       piiH-i'ss;.       ,'|1|.|       ull.'ll       .'N.-.'SSIN. 

.piaiiliti.-s  ..t'  a.'i.l  iiialM-  ilinr  app.-ar.iiii-.'  in  tl i-i.'aiii>iii,  tin-  li\.-.|  alkali 

iiia.v    .,Mt   l„.  .,iitli.-i.-iit   t.i  ii.'iiti-ali/.-  i!  Jill.  S.I  that   aim ia.  ili-i-i\.-.l   fmiii 

till-    l,r.-;ik.|..VMi    lit'    aiiiiii..    .i.-iiU    .  jiai.'.-    till;-.     iiist.-,i.|    i.T    li.-iii.,'    .-.iin.Tl.'.l 
ill'"    III-. -a    is    .-inpl.pw'.l    tl.    iH'iiliali/.'    th.'    .-n.  .'ss    ..i"   aci.l.       M.ist    «i.|-ki'i 
haM-  ill   this  \\,iy  i  n  p|;i  m,.,!   ,i,,,   ^,.^.^    |, ,„,_,,,  .•|,iiiiiiiiii.-i   cxim-.-I  l.iii    th.-il    h,-i- 

hiiiL'  li.Tii   kn.iwii   t. .-lii-  ill  vii.-h  .•iiiiili!iiiiis  as  .iialii-lii-  a.-iilnsls       Smii. 

l-.'i-.-nt     \M.|-K.|-s    ill-,-,    li..\.  .'M'l',     iii,-iiii,..|    In    .pii'sti.iii     the    si._'iiilirail<'r    III 


AniH>sis 


iiiiiiiiiia  In  tliis  riiiincction.  Iii'lii>viii<^'  tliiit  llic  ini-iciisi'il  aiiiiiKinia  cx- 
.lidii  is,  lil;i'  llu"  acftiilic  luidii's  tliciiisch  i-s,  a  iiniiluct  of  |U'i-\crttMl 
.tMli"liMii.  M>'  tills  as  If  may,  it  is  no  doiilit  tiiH'  that  aniiin>Mla  Is  uscil 
.!■  ii''iitiali/iii'_'  ai'iij   In  tliscasc.   altli(nii;li   It    may   not   lir   an    Ini|njr1ant 

H'liir  in   llic  malntciiaiH t'  milt lallty  iimltT  nmnial  comlilioiis.     It   is 

I'm. till'  ot  sal'i'ty.  In  tliat   It  liclps  to  caic  fur  an  Inrrcasc  in  itcl'l  wlicn 


iIk'  iiiiiriial 


nil  'lianisiii  i>f  the  IichIv   Is  o^ci'taxcil. 


Kxrifiriiiv  (iK  riinsniAi'i  s.     Tin-  III"!!'  pninancnt  ciiiitiu!  nt'  ncntralit.v 
!■],  mis  nil   till'  cMiitliin    cif  |ilios|iliat('s  hy    tlic   klilm-x.      Tlic    iirinclplt' 


\i'iniiiir  tliis   proci'ss   is  cxactl.v    tin'  same   as 

innitliiii    \\itli   caiiiiiiiir   ai-iil       In    tlir  nut'  case    it    Is   tl 


that  alrcadv  fliscnsscil 


11'   \iilatlh'  ai'nl 

'  '»,  ami  In  li thiT,  the  fi\<'ii  |)hi)N|>hiiiii'  acid  that   Is  (•(iiicci'iit'd  in  tin' 

i.,iitinn.     The  latlii  IpcIwccii  tlii'  arid  salts  <<(  i(h<is|ih<iiic  ai'id,  MIL.I'O,. 

.iiid  the  alkaliiii'  ^Its,  .\1.lll'l»,.  In  lil I  is  a|)|ini\iiiiati'ly    I   tn  ."i.  iiiit   In 

ill'  iiiint'  thin  latin  varirs  ai'c-cudiiiir  in  tin'  amniiiit  id'  II  inii  that  must 
!'<'  I  limlnati'il  I'miii  the  Idnnd.  In  ntlii'r  \(  nrds.  ,i  di'linltc  amnuiit  nf  jilms- 
piinrii'  acid  i^  i-naldcd  tn  rairy  xarialdc  .imniiiils  nl"  II  inn  nut  nl'  tiu'  Imdv 
li\  I'aiisiim:  till'  aiiinmil  ><(  alkali  <xi'ii't.'d  in  I'nmliinatinn  with  It  tn  he 
.  MiiM  altt'i't'd.  i''nr  cxampli',  in  the  I'ni-m  id'  MM  I'O,  a  iriwii  amnunt  "!' 
I'll  nln  iniisi\  i-:rrirs  nut  iimri'  ||  inn  thai,  w  lii'ii  it  i--  i'mtcIi'iI  as 
\|  lll'li,      Till'  ad.iustmi'iit    lii'twci'ii  thi'sc  1  w  n  s;ilts  is  ;\   tiiin'tinii  nt'  tln' 


W  r  nia\    aci'ni'diiiLdv    iiii'asuri'  tlir  ainnunt  of 


ali  ri'lalii 'd  '>v 


iiranism   1'^    findintr  Imu    mni'h   slaiidaidl/cd    alkali   must    lu'  addi-i 


tl.  M  i.'iM'n  <|iia 


ntitv  id'  ni'iiii'  until  tlu'  ii'artinn  id'  the  hlnml   is  nlitiunt'd 


till'  latter  valiii'   is  I'nnstaiit,   the  titratinii   ran   lie  diiin'  simply 


tialini:  till'  nriiii'   willi   an    iiidn'atnr  siu'h    a> 
it   at    aliniit    I'll   nl'  Idnnd 


siilphiiMi'phi'nnJplil  I 


hi.|i  I  li;iiii:is  tint  at   aliniit    I'l,  nl   lilnml 
A   iiinri'  si'r\  ii'i'alili'   indiratnr  tn     isi'.  imu  I'v.-r.  i^  plinui};tlith,iti  m,   ln' 
iii^c  its  rnd   point    is  siii-ii   that    w  In'ii   hinmiii   iiiiiir  just    n-ai-'v    in'iitral 
II      that    is.  wlii'ii   till'  titi'alili'  ai'ld  approach. -s  /.mn     thr   i  ( i    .disorl.- 
■1  [iLwi'i'  nf  till'  i>lasm.i   i>  !it  its  ma\lmum  ol'  >«>  \  nU    pi'i-  i-i'iil   and  tli»' 
iii.iiii.i  I'XiTi'linii  hy  till'  iiriiii'  is  ziTii  I  \'an  Sl\ki".      It  i>  .nh  .intatri'nus. 
•  11  inii',  to  iisi'  this  iiidii-atnr,  hi'i-ausi'  it    haiipt'iis  tn   hjiM'   its   tnrnini: 
;'i:   ^itiiilt'd   tor  n   ri'artinii   « hirh    is  «i'll    to  tin'  alkaiiiii'  sidi'  nf  ui-ii 
".  .iiid  mIiIi'Ii  is  icai'hi'd  in  nrinr  wIu'M  the  lihod  i-  a(    its  m.'ixinial 
.iidiininjr  powi'f  and   im  ammonia   is  ln'inir  us.'d    Inr  iiiMitr.ili/atinn 
-1-^       As   till'   ( '( '    .'ondiiiiiiu.'    poui'i-   ot   Ihr    lijnod    driTi'tisi's     thi'ii 
!    tlii'i'i't'ori'.   Ill'   a    lu'opoi'tionatc    iiirri'asi'    in    ammonia    and    in    tin 
!■■  ai'iditx    id'  till'  nrinr. 

niiu'li  a  irriii'ral  piiralUlism  i-xists  'irtui'i'ii  tlii-.'  \;iiiirs  in  r.is.  >  id' 
'  ^.  I'll'.,  thi'l'i'  is  nn  sirirl  propolt  lolialitv  ,  Tin'  rvpidirni  has 
'I'-  liri'U  trii'd  id"  comparintr  thi'  alkaline  ri'siT\i'  ot  lln'  hlood  uitli 


,j.<: 


vsBsss;jsagaaitsxsn 


4S 


I'liYSKdi  lu'MKAi.  iiAsis  or  piiYsi()i.(Hii<\i.  rRO«'i:ssK>? 


the  cxciPtioii  rjite  of  acid  as  (Ictcriiiiiu'il  ))v  an  application  of  Ambanl's 
ciiuation  for  chlorides  and  urea,  and  witli  ciiriouslv  sjitislactory  rcsidt^ 
^Kit/and  \  an  Sl.xki'K     Tills  c(|iiation  is: 

r.liiod  coi titration       constant  \  ^  y^  ^(';   where    I)   is   the   excretion 

rate,   \V   the   l.od.v   wciirht.  an<!  C   Ih. iicentration   of  excretory    i)rn,l 

net  ill  the  urine.  For  the  pres.-nt  pur|)ose  1)  is  tlierefore  the  n\inil>er  m 
e.c.  of  N  1(1  alkali  lor  acid  i  reciuired  to  l.rin^'  the  urine  to  the  neutra' 
point  of  pheii.ilplitlialcin  i>his  the  N'll  ,  expressed  as  c.c.  id"  an  N  10  solution, 
for  the  twenty-four  hours,  and  ('  is  ce.  of  N  10  alkali  and  of  N  10  Ml 
per  liter  of  urine.  If  we  assninc  that  the  acid  accumulation  in  the  hloo.l 
is  iiroportional  to  the  fall  of  the  plasma  CO,  fitrnre  hclo'V  the  maxima- 
(ijiiire  <d'  SO,  the  ahove  eciuatioii  liecomes: 

l> 

Uetained    acid       SO     i)lasiiia  t'O.       constant   x  v  \V  ^'  • 

For  practical   purposes  it   is  liest   to  make  the  necessary  analysis  on  .i 
sample  <d'  urine  collected  over  a  period  of  one  to  four  hours,  an.l  to  col 
leet   the  lilood   for  determination  of  its  reserve  alkalinity   in   the  middl 
of  this  period.    The  lu  .nt)  four  hour  rate  of  cNcrelion  is  then  compnlr- 
( I)  I   from  the  anal.v  sis 

The  value  calculated  l>y  the  alxive  ei|uation  has  lieen  foinnl  to  auMc 
with  that  of  the  (•(l.comliinintr  powi-r  of  the  plasma  to  within  10  vo 
unies  i>er  cent,  except  when  l.icarlioiiate  is  lieinu'  taUi-ii  l>y  the  persoi 
when  the  IiIoihI  liicarlmiiate  is  much  liit;hcr  than  in.licated  liy  the  iirin 

:!.  Determination  of  Alkali  Retention.     Another  valuaMe  criterion 
the  alkaline  rcs<r\c  is  t|i.>  amount  of  alkali  re,|niied  to  eliam;i'  the  i 
action    of    the    urine.       In    health    the    « '„    of    the    urine     varies     fn... 
(i(«M),(tli;  .V  I  !•„       4 >i  to  al.oiit  0.1MMI.0(MM>:!.">  A  i  1',,       7.4<i)  with  a  no, 
of  alioiit   0.0(M),(Ml|   .V   I  r,i       (li.     These  rxlremes  are  rarely  o\erslcp|.. 
in  disease,  lnif   fiti|iienl  ly   the  nrinnii    is  consii|eralil.\    ditTerent.      In  im 
dio  renal  disease,   for  example,   the   iiie:iti   acidity   may   he  .ipproxiiiiat. 
ll.dlHl.OOri  .V  I  I'll       .".  .;   ,  or  live  times  the   normal   value.      A   ei-rtain   ■! 
•_'ree  of  acidosis  is  lli.-refoie  eomiiioii   enoii-rh    ill   this  condition     a    t     ' 
which  has  iiKlicated   tl.*"  advisal>ility  of  administcriim:  sodium   liicail" 
ale.     It  has  lieeii   foiin.!  ihal  .'i  '_'rams  or  less  of  so.la.  ttiven  hy  monlli     ■ 
a    normal    person,  caiis.-s  a   distinct   diminution    in    the   ( 'm    of  the   nn 
whereas  in  patholouic  eases  it    may  he  neccssar.\    to  i.'ivi'  more  than    '    ' 
i;rams  I.efore  a  similar  effect   is  oliserve.l  (  L.  .1    Henderson  and   {'aim 
aii'l   Sellar.ls"!. 

{•"or  this  very  larirc  lioldinvi  hack  of  alkali,  the  oriranism  and  not 
kidiii'v  is  ri'spoiisil.ie  This  is  indieati-d  liy  the  fai-t  that,  when 
administration   of  alkali   is  diseontiniled,   the  a<'idity   of   the   urine   -      ' 


ACIIXISIS 


4!) 


nuiiins  its  olil  level,  iilll U'li   iniw   if  ;i  Miiiillcr  ilose  iif  alUiili   is  '.'iveii, 

till    (',1  «<  tin-  iiiiiif  will   iliimnliiitely   lie  liiw.rcil.     Tlii-sc   t'ii'Ms  inilii-jite 

III, it    lor  till'  ilniilc  (iefirecs  ci'  in-iilcisis  iMiiimioti   in  rlin.iiic  iIimms.-.  tin- 

|>ii«i>i'rly  (•(iiitinili'ij  injiniiiisli-.'ition  u 


I   siichi   IS  \  I'I'V   lll'lili.'lli 


Iv  :i  most  ;iil\;in- 


l:iu(iiiis  liciitiiii'iit. 


^i-iV  iivif  \tsmaasi0t. 


CIIAI'TKI!   Ml 

('(•I.LulDS 

Siilpst;inc(s  uhirli  .•.•m  lie  i<l>t:iiiii'<l  in  tlic  iTystalliiic  stjitc  iiiiil  wliirli, 
wli'ii  in  snhitiuir.  jiic  (•.•i|i;ilili'  „(  r.'jidily  ilitTilsiii^  llir'oimli  nifiiil)l';iiics, 
iirc  (Irviv'ti.ili'il  iis  civ^tjilli.ids,  iiiiil  iiiT  to  111-  (lisliiiu'iiislii'il  I'lmii  iimitlicf. 
l.iiiT,,,-  ir|-,,ii|»  III'  siili-.t,iiii-i>s  nut  li.-ivinir  IIh'sc  clijifiifli'riviii-s  nr  liiixini; 
tliirii  iihly  in  Miy  rninnr  tli-_'i<i'  llif  i-.illiiids.  In  I'vciy  liclil  of  i-licin 
i>lr\  tlir  |iiii|ii'itii'S  ot'  rolloiils  li;i\c  iircn  stUilicil  cNlrusix  cly  <lll|-ini..' 
nrint    MMis,  Inil   in  no  lirlil  more  tiiiin   in  tliiil   wliii'li  <'ovfrs  tlic  clicrii 

islry    lit    liiii|in;ii-;il    lliijils    iinil    ti>.Niii's,    into    «  Iiom Mi|iiiNilion    rolloids 

inltr  nini'li  nioic  cxtnisivily  tliiin  iMystnlloids.  Tin-  Miliji-rt  of  folloid.il 
rlniiiislrv  li;i>  ihdii'd  ln'i-i  mr  so  rxtrlisix  f  tliiil  :in  iittcnipt  to  <|o  iiini'' 
lliiin    indiriili'   som.'    of   ilo'    most    inipoitiiiit    (diiiiartiTistics    of   rolloids 

vMMild   Ijilvc  lis  t';ii-  lic.Miiid   till'  liniil.'ilioiis  of  tliis  I I<,      Tllf   f:il-|-r;i(dlin-.' 

■  ilil'lii'iiiiiiiis  of  ilic  snlijci-l   ill   |ili\si(i|i  jry  ;iiid   iiifdirinc  iirc  only   lic^jn 

lllim    to    111'    I'l-.lli/ril. 

'I'lii-  tiiiii  ••lulloiil,"  111  '•riiljiiiij.il,"  dors  not  rcfi'i-  to  n  cljiss  of  i'lii'Miii-;il 
siilist.ini'i's,  liiii    ijillii'i-  to  :i  sliilr  of  ni.-ittir   u  liiidi   is  i|uiti-   iiid<'|ii'iidriit 

of  till'   I'liiiiiii-iil    .I'liiimsiiii  n   iif   till'   siilisi; W,.   jii'i.    fniiiilijii'   uitj^ 

nil. re  colloids  in  tic  orii;inii-  th.-m  in  t||,.  inoiiriinic  world,  yet  tiii'\-  iin 
plriitifnl  ih  liiiiji.  .'inil  tlic  miiiic  snlistiiiirc  iiuiy  at  one  time  lie  colloiihi! 
•""'    ''  'I'liplliir  iii'iii'uJlniilMl.        Indi'iil.  iiiidiT  iifipi-opiiati'  conditions  pi-nl' 

alii      all  siilistaiiccs   may   assiiiiic  tl Iloidal    state      not   solids  and    lii| 

'"         iloiic,  lint    LMsi's  as   Will.      It    is  mainly   with   liijnids,  liowcvcr,   lliai 
1-  ciiiici'tiii'd   ill    liiiiclicmisti-\-. 


CHARACTERISTIC  PROPERTIES 

'I'll''  di-ilii,  linii  liitiMiii  iiiiilii-nlai*  and  <'ol|oidal  solutions  is  a  rcla 
li\i'  line  Siipp.:-.i.  tiir  I'vampli'.  lliat  \\r  take  a  piece  <if  ^rold  in  uatc 
all. I  iii\ii|r  it  lip  iiiln  siiiallir  and  smaller  parts.  At  a  i-ei'tain  s|ai;e.  lli' 
pallid'"-  uill  I  e  Ml  line  lliat  ll;i'>  \\\l\  ninain  in  snspension  and  lie  in 
\isilile  hy  iir.|iiiar\  means.  'I'liev  aie  then  said  to  lie  in  the  eoliuiil; 
^talr.  ir  We  ili\iili'  tliiiii  liiilhir  until  tliey  lieenine  mnleenh's  of  iri'l' 
a   nioleciilar  siilnlion   «  ill   I Iitained.      In   the  colluidal   state,  there  a' 


■  \l-.l..  iil..r    ...I,  n...,-     ,,,.  I,. I.      il,,. 
ri.Mtii.     v.ilK. 


;..  i...i,i...    -:,.] 


t.iT.     u.,|    ,l,,:i..KTt, 


nii.i.dins 


il 


H 

i=3 


1 


two  (listiiii-t   pliiiNi-s  iti  till'  snlutioii,  iiiK'  solid  iiiiil  t1 tliiT  lii|iii<l,  iind 

l.riwcrii  till'  two,  lii'i-aiiNc  of  tlic  irii'iit   MilHlivisioii  of  iIm-  orisriniil   I'lif- 

lii'li',   is  iin   ciioiiiiolls  sllll.K I'  i-oiiliirl.      'riic   solution    is   III  II  riii/i  III  mis, 

:iiiil  at  the  inlriracc  lictwctn  tlic  two  "  pliasfs"  the  plivsical  I'oiws  wliicli 
.|i|ii'Mcl  on  smt'.icf  c.  (.'..  SIM  rare  Icnsioii  I  si'f  jiasri'  (>4  i  an'  I'liiu'iiioiislv 
,|i\clo|)<'il.  anil  ail'  nspoiisiMo  I'or  tlir  piniliar  pnipi'i'tirs  of  i-oljoiilal 
solutions  as  i-oiiipaiii|    willi    tiiosc   of   molfriilai'   solutions,   wliii'ii    iiiav , 


'ii'i'ori'.  III'  st\lril  iiiiiiiiiili  m  mis. 


'I'lic  siiliitions  of  ri'vslaliini'  siilistancrs 


whii'li  wi' 


liavi'  liitlirilo  l)c('ii  I'onci'rnril   witli,  aii'  lioniom'in'oiis. 


r.rtwi'i'ii  tlirsi'  luo  (■loiips  of  solutions  is  all  intciiiiriliati'  our  nanu'ly. 
^iisjii  iisioiis  las  susprnsioiis  of  i|Uart/.  or  i'::iliou,  or'  oil  I'lnulsioiis ).  I'c- 
-iilrs  li.'ini:  tuiliicj  in  transinittcil  liiilit,  tin'  solutions  may  l>i'  situ  liy 
means  of  the  iiltiaiiiicroscopc'  to  rontaiii  pjiitirlcs.  Tlicsi'  can  lir  si'pn- 
riliil  liy  (iltration  from  tlic  lluiil  tlicy  arc  suspciiilc.l  in.  except  in  the 
case  of  many  emulsions  in  u  liidi  the  particles  can  si|ueeze  llicir  \\  ay 
tliroii'_'li  tlic  liltcr  pores  li\  clian^jiiiL'  tlieir  sliapc.  (In  staiiilins.'  or  liciiiir 
I  ciii  lifu'.'cil  siispeiisions  max  also  separate  into  tlu'ir  eoiistitiiints,  al- 
ihouirli  tliis  call  lie  irreatlx  liimlcreil  liy  tlic  adililion  of  a  siispciiiliii'-' 
sulistancc  siicli  as  nelatin  or  eeitiin  lio<lies  liavinc;  a  so  called  protcc 
li\c  action    las   peptone,   proteose,  etc.). 

True  Colloidal  Solutions 

1    The  Solution  Is  More  or  Less  Turbid,     l-'icpicntly  this  .an  l.c  n i: 

i.i/e.l  liy  holding:  the  solution  in  a  thin  walled  irlass  vessel  ai:aiiist  a 
■  l;irk  liackirfoiiiid,  luif  the  tiirl.idity  may  he  so  slit.dit  lliat  it  reipliies 
lor  its  d.'tectioii  the  Use  of  tlic  Tvndall  phcnoiiicnou.  This  is  familiar 
!o  all  ill  the  elTect  of  a  lieam  of  siiiili<iht  let  in  ihroui;!!  a  small  aiiertiirc 
into  an  otherwise  darkened  room.  In  the  course  of  the  lieani  suspended 
•  lii^l  particli's,  which  arc  invisihlc  in  an  cipiall.x  illuminate. 1  ro.'iii.  1m- 
•111.'  \isilile.  an. I   thus   ren.lcr   \  I'l  .\    .listin.'t    the   pathway   of  tin'  beam. 

I;  ; lloiilal  solnli.Hi  contain. '.I  in  a  '_'las>  v.ss.'l.  prcf.'raM.v    with  paral 

'  vid.s.   i»  held   in   the  course  ..f  such   a   h.'am.  lln    Tiiinliill  jihi  iimiii  iimi 

!  I..' s. •I'll  ill  the  lii|ui.l,  whi.'h  is  not   the  cas.'  with  m.il.'.-ular  soluti.nis. 

iw.il    artifii-ial    liudit    ma.\     he    emphi.\.'.|    tor    int.'iisifyini;    the    clTc.'l 

'    ■    liuht  that  is  s.'iit  out   at   riyht  aiiul.'s  to  th.'  Iicaiii  is  plan.'  p.ilaii/i'.l. 

•Il  111. 'alls  that  the  ])articles  retle.-tiim:  the  liirhl   iniist   lie  siii;dl.r  than 

111.  an  w  a\ .'  l.'iiirth  of  tin'  liiiht  f.'riiiinir  th.-  Ii.'aiii,     It  sIduIiI  Ih'  nii.l.'r 

■1    that    the    iii.li\  i.liial    partich's    tli.'iiis.'hi's    iiiav     n..t    h.-    r.'ii.l.'r.'.l 

^■l'-  III  the  n.-ik.'.l   .'.\i'  li\    tli<>  heam,  alth.niLrh   in   such   cas.s  th.'\    .-an 

••  !i  11.'  sciii  hy  usin.j:  int. use  illiiminati.in  and  a  .lark  li.l.l  i  'illraniicr.i 

'  ■■     .■.'iiiliin.'.l   with   snilalile   mairiiiti.-ation    i  l-'i'j     111'. 

J   Colloids  Do  Not  Readily  Diffuse.     To  .h'l istr.it.-  this,  t.'>i  tuhes 


.»_  I  in-liMi  Ml  MM  \i     i.\^i»  111    i'ii\ -iiii  iH.ii  M    riiiH  rssi's 

.III'  ll.lll  lilll.l  \\llll  :i  ')  |M'|  rI'Ml  ^cillllliill  nl  (tlllr  'jrlillill  n|'  a  I  |icr  c-clit 
^I'lllllnll      kI       {iIIM        .P'MI,      .IImI,      .lllrr      ill'-      .jl'II.V        !>•      M'l,      till'      slillllillll      lllnlll 

I  \.itiiiii;il  mil     1^    |Hiiiii.|    Mil     tlir    ^iiihii'i'.    Ill,     w  lull     il     I-.    Ill     Inuli     ^|ll■ 

i-llh  'JUnir  llir  llllir  nl  •^^ll.lllll,  I'll',  1>  |llill'i'll  lllnlllll  I  |l  IW  1 1  W  il  11  l>  ill 
llll-    Millllliili  III     I  III      I'ilsi'     III     I'lilliinl.ll     ■^ll|lllill|l^     MTV     lilllc     ir    illlV      llilVll 

■«MiM    mill  llir  L'l'l.il  111  111    ;r^:ir  uill  nri'iir.  ('\i'ii  iilliT  si'MTji)  (|;i\s;  W  1icIi',i> 

lllll'     illnli  illlal     viillllmils    Hill    ihlVlI'M'     llll     il     I'lillsilji'l'illill'    llistillKM'.        WIh'M 

riiliiii'ij    ^njiilmii-.    iUT    iisi'ij.    llll'   i|irfii>.iiiii    ciiii    ii'ihlily    lie    rci'iitrni/i'il    li> 

VIMIill  Ml--|lll'l  lull  M'l'  |-'|ij  l.'l  ,  lllll  H  lull  llll'\  iiri'  I'llllllll'SS,  till'  |iri'S('l|ri' 
nr  ,|liM  III-.'  nl  iJllTll^li'll  lllll^!  Ill-  iji'li'l  IIIIMi'lj  li\  li'llli  l\  ill'^  tllc  rullllllll 
111  '.'ililllll  111  .I'.MI  ilMil  i|l\|i|lll'J  il  illlii  ■«l|i'('S  111'  i'i|llill  >i/c,  \\  llii'll  lire 
llli'll    I'NillM  :lir'l    I'lli'lllii-illlv      llll     llll'    ".lllislill Ill    i{Mi's|  lull. 

A  hiiilii'i  II  si  i^  .'itViii'ili'il  li>  llll'  liiiliiiT  111'  I'lilliiiijs  til  ililTiiM'  tlinmuli 
iiH'liilir.'ilii'N  iJi.iUsis  This  w  il',  Ihi'  iiH'lliiiil  nri'^iiiiiillv  ilvnl  liy  'riliilMli^ 
'iiiiliiiMi  In  'livi  ih'^iiisli  liilwi'iii  Miiiji'i'iiliir  iiinl  I'lilliiiiliil  Miliitiiins.  Tin' 
-iiliilmii    iimli  r    <'\iiiiiiiiii1  imi    i»    Mliii'cij    in    n    iliiihrii;    wliirli    !•<    tlicii    iiii 

IMiTni'iI     III     il     Hlili'     Vi'ssil    I'nlilililliti'j     llic     |ll|li'    >.iil\i'llt  Till'    nlijlT     t'lU'lll^ 


1^1-         ;■.•,■:•-:'■!■'■>:.  ■     '        ^.      '      „■"     ,'.     '       .1     ...:i     |.;,,1     ..!■. .„.,'-         Tlir     Illl.,.^;-. 

nil  II!       I    .h.i:  l.i.ii."    •,    It'    .    I,      ll.l-     I-ir-     ni.i..v,.    Ill,     ,,l,...i  |.ln ,      ,11,,  I.    ■!     Ih,      -■irl.i.,-    ,.t     III,      I'l.i 
■"  --     '     '•■      -'.    '.  .-..I     1         .       ■   I.M'.      Il    '       ,,,!!.    .'!.,;      ,    ,l:,l',,i    . 

"I     lilill.Wil      1 -iviiil     in     '.;i|iili||     nl     il     lii'll    s|lil|li'l|     m|;|ss     \  I'ssi'j     I'jlls.'d     In 

111"     "ilii     |iiilrliliii'lit     I'iil'i'i',     lllll     iimi'i'     I  I'l'i'iit  l_\     sii  i-iilli'ij    ijilTllsiiin    Sill". 

Ilil\.'      I"  111      , 111,, (lllll  'riii'vi-      I'lilisisl      nl       |lii:     HI-     lisll      lljilljllcls     III'     III'     I'lil 

l'"l sill's  'llll'     Lilli'l      ill','      lllillji'     ll\       |ilill'ill'J     SIIMH'     I'lllliiilillM     llisMlJM'il 

ill  lllll  1  III  ;i  t''st  liilii'.  wliiili  is  tliin  nlli'ij  sii  iliiii  till'  I'liljiiiljiiii  niiis 
"III  ''M'.'lil  Ini'  il  tlilii  linii  \\  llii'll  ii'iiiiiiiis  iiijlii'i'i'ii!  Ill  Ihr  Willis.  Wlli'i' 
'III'  I'nilniliiiii    liiis  si  I.  ill,    ^.11'  I'.iii   111'  ri'iiiii\i'i|  iilli'i'  Iniisi'iiilii;   It    li\-  iiijiiv, 

ill'2    il     lltlli'    Will, 'I     In    lln\(      |.i'|\M,.|i     ill,.    ^,'||-    ,'|ni|     llll'    Hiills    ill'    till'    trs|     lllll. 

'I'll''      --il'-      tillllih'J       llll-      I   iillnhl.ll      snhlll'.l!       is      llii'll      si  Is  |  Ic  I II  j  I'l  |       ill      Hilllt 

"I       snlili'      nl'      llll'      s,,|\,'|||       II,,', I       ill       lili'l'ill  illU      till'      rillllliljilj      sllllltillll,      l-il' 

liriii'.:   liik.'ii   lliiit    till'   llll  iiisi'i   111'  till,   lliiiils   iiisiiji'  ami   nutsitlc  ut'  the  s,j. 

s'illil     ill      'In-     s.lllll'      I.'\i'l.  .'''<n|ili.I  illil   s,     |.s| iil||\       W   ||,.||     I'llllllljjlill      Sill's     ill' 

lis,., I.     s,,||H'     I'liMnlll      lll,l\       ill       lilsl      llllVlisi'     t  |l|'ill|'_'|l.      lllll       ir     till'     lllltl'l'     lllli' 
!|||1'      l|lill\si,ll  is       I'l    l,,'\'.  i-il      iihij       llll.      i|ii||\sis      illlliwcil       |(,      IM'tll'I'l'il,      II:. 

riilsis 


l^"! 


1  Ml  I  nllis 


5.1 


Wlicii  ii  lliiiil  Mihitiiiii  cxliiliit^  linili  111'  ilii'  mIiiim'  |irit|(filirN  i  i  c  .  tin- 

T.lnlilll    |illi'll(iliiilliin   illlil   illililVu^ili|lil>     .   till  II    liiM   lir   111!  ilmilit    as   in   its 

I    a-,   riiiiijii 
I   IMM\    i-\hlliil    a    \iT\ 


Imiiii.'  ill   II   'II' Iliiiilal   slalr,   Init    llnti'  ai-'   >i|listaiii-«>.   mm-I 


ml  III    |i|iili'iii  siilntiiiii 


il'   rrllaill    »l  nii'.'l 'is.    "  liiil 


sliu'llt    llilVusillilil>     ill    :l    ilillwiT    llllt     IM'I     -I   ■     ■      !  In      T.MIllall     |llll'l|l>llll'l|llll 

Siilistaiii'fs  lit'  this  '.'lull  1 1  riiii>l  ililli'  ti  !ii  a  I   I '  |m  s  In  I  w  ri  li   iiinli  i-illai 

.itnl   nilliiiilal   Miluliiiiis,   ami    In   iIi'Iimm,  ii-    liiir    nalllli'    II    is   ihtis 


i 

n 

i 


-l)'»     .|ill-]-...n     p  ,,.    ^,  l,,,|.>     ,.1      , 
...II. .1,1     -l,,:>       I  ,     „ 


.i-i.,:i....l    -I..:. I    r.    .'    .1-1    Ii..    it"i..lin...-i-.i!    ■•!    i 

,  I   •    V      U        I  I.Mv.ll  I    ' 


I"  .:ii|'l.i>     1. ■line. I    iiii-tlLils   siii-li    .1-.    ill...,     ..1    iili  r.iiiii.-i  ..si'"|i\  .    iillr.i 

■■•ill"M.    I'll-,    wllil'll    ran    lint     111'    ili'siM  iliril    liiM->.. 

3.  The  Size  of  Colloidal  Particles.     Ii  mil  I-  aiipai-ni  iliai  ili.- .-..ntial 

i  •  '  i>    n|".n    \>  liii-li    I  111.  alinv  r  im  nl  inn.,!    |.|h  ii.nii.  ii.i   .1.  pi  n.l    is   lli.'  -1.  '■ 

i '■  |i.iilii-l|.       I'arlirlis   w  liii-li    .•.111    still    lir   siin    iiinlir   iIh-   niii-i  ns.-np.' 

'•  I'li'rj    inirriiiis       '\'\\\-\    liavi-    l.ii-n    .  ..iii|iiit..i|    In   liave    .i    iliiiniisinii    ..l 

'Mllll    liiiii.      nf  iiiiii-c.  ami   llir.\    I'M  Ml   si|s|iinsiniis,      I'aitii'li's   \' hirli 

■isilili'  iiiiriiisi'n|)ii-,ill\     uii'li-r   till     nii|iiiai\    I'nii'lil  imis    nl'    illllliiiiia 


.>4  niVMi  ut  Ml  MUM,    |i\-.|s   OK    I  IIVsiiil  iH;)r\l,    IKiMlssis 

IIMli.      Iiiit      nil'      slill      \  imIiIi       t\  i  I'll      till'      lllli   i:illrliiM-ii|iii-      JllmilillMliiiM      Iv 

UM'il,  III iilli'il  Miliiiiii  runs.     'I'lu'V  li.'ivi-  !i  itniiriioiiiii  lii't"''!'!!  Ill   /(  mill 

1  fin  ilt(HMI,(HI|   Mini.  I,*  iiliil  llicv  I'uiisl  iliili-  till'  nilliiiils       riirt  ii'li',  miiiiIIiT 

lliaii   I  /i/i  ;iii'  i'.illi-<l  !iMiii-iiii'<>.  tlii<>  ti'i'iii  Ix'iiiu'  u» 'I  III  iiii-liidi'  t.iii-  mill 
(■ciili'H  iiiiij  ii>ii><  piTM  III   ill  iiiiili'i'iiliir  Miliitinii^      I 'I  III'  iiiiiiri'iiii  III'  hyili'ii- 
iri'ti  is.  I'lti'  i'\iiiii|>l<'.  riimpuii'il  til  III'  O.titiT  In  II  :">''  fifi,  and  that  ol'  «ati'r 
v,'i|iiir.  II  Ij;!  /i/i  i      'I'liis  I'liixviliiMtimi    lit'   iJissiiK  I  .j   siilistalii'i's  ilri'iiidjiitr 
til  tlir  si/r  III'  till'    |iiil1ii'li       .'Hill    liiii|irul(>   ^l|ll^^  ^    lllf    li'lat  I'Misllip   iif  iilii' 


l'l,|>s    nl'    xI|I,n!;i||ii-,    111    nllli'ls  All     iili.l     111'    llli      |-|'l,l!i\l'    si/cs    iif    I'cillliiil. 

IMi'lirlr--   ;iiiil    mull  '  .ili'>       I    riiiii|i;in'Min    witli    •<iii-|i    raliiili;ir   nliji'i'ts    ;i-- 

I'l I    I'lil  |il|Ni-l,     ;iiii|    ,111    ;i|illiln\     liai'illll>    1-    L'iMIl    III     I'm      H        Till'    till 

ill    wliii-li    till-       |i,iilii-,.     "    is  >iis|..  liijiii    In  I'.iM.ij    till'   ili\f)i  rsiiiii    iiif  ihiiiii,  < 
iilifihil   /(/m/v,  ,  ,'iiul    lii,'    imrtii'li-   ii>i'li'  iIm    ili^/ii  rsoiil.  ur   iiilinml  plni 
It    i>  till'  iiim  iiiiiii>  i|i'\  i'lii|iiiii'iit   III'  Mil  iih'i'   \vliir|i   di'ti'i'iiiint's  llu'  il- 

*«  """I     liiTi:  .    aril    uo  nmil'.l.nl     ,•  ., 


(OI.MMIW 


65 


ri't'i-iirc  ill  till-  |)rii|ifi'tif.s  III'  ii  riilliiiiliil  Miliilimi  I'rniii  tliiiNc  of  a  ><ii>|M'ii- 
-.loii   III'   till'  siiiiH'  siilislaiii-t'.     TliiiH.   tli<>  ililTi'n'nri-   IicIwitii   a   cnlliiiil.il 
siihitiiiii  III'  pliititiiiiii   I  |ii'c|iat'i'il  liy  allnwiiitt  iiii  rli-i-tri)*  iiir  tn   furiii  Itf 
Mm'I'Ii   |iliitiiiiiiii   cli'iM null's   ill   watft'l    ami   piriM's  uf   pliitiiiiiiii    in    wati'i' 
.|r|it'tM|s  (III  till-  fact  tliiil   lilt'  Ml   t'ai'i"  III   tin-  |i|iiliiniiii  in  llii'  fmiin'r  ciisc 

li;is  I II  iiiri'i'iiM'il  niiin.v    riiiHiiiii    I  inns.     Winn  tlit'  snliiiiv  isjuii  In iik-s 

^tiij  (.'iralrr  anil  llic  pai'ticli's  cain  tin'  si/.c  iit'  nmlri'iilrs,  tin-  |ilii'iiiiiiii-iiii 
ijili'  tn  sllifai'i-  <ll'V('ln|>iii('llt  liiTiiliM'  .sllpprt'sscil  ami  tlmsr  illlc  In  cull 
ri'iiliiitinii  ill  unit   Miliiii'i'  Im'ciiiiic  accMitnatril.     Tlic  prnpcrtics  tlcpcinl 

rllt  nil  iisMinlic  prcsslll'i',  ililTllsiliililV.  etc.,  arc  i'\lliliili'i|  li\    ail  ilispclsniilH, 

vvIii'IIm'I'  inn^.  iiiiili'i'iili's  nr  piiiticlfs,  liiit  siiiiii'  iif  llii'sc  pi'iipi'i'tii's  arc 
iiiiii'li  liiiil'c  pinininnccil  wlicii  llic  liispi'isniils  arc  nl'  lar'.'c  iliiiii'iisiinis 
Mini  iillicrs  \\licii  tlicv  arc  Miiall.  In  ntlicr  ^^<l|■^|s,  tlic  plicnnincna  iliic  t" 
«url':iri',  siii'li  a.s  tlms"  nf  .suif;nc  tciisinn  (SCI  pai;i'  lilt,  Ihiiimii'  Miiparcnl 
iiiiiv  wlicii  the  ilispt'i'siiiils  liavc  I'lic  prnpcrtics  nf  matter  in  mass;  xilicn 
llic  ilispiTsniils  liccmiic  miilcciilar  in  si/.c,  tlicV  manifest  llic  pnipcrtics 
(•li.iiactcrislic  nf  line  snliitinns. 

4.  Electric  Properties  of  Colloidi.     Must  cnllniils  carry  a  dun '.'<',  which 

iii,i>    Im'  cither  piisili\c  III-  ncuatixe  tnuaiil  the  ilispcrsiini  incilinni.     I'lnth 

1  (•>  stailiiiilx  aiiij  eiijlniils  tlicrcfiiic  cany  clcetric  char'.'cs:  in  the   t'lniiicr 

•  ■'SI',  liii\M'\cr,  the  chaiire  iliics  imt    reveal    its. If  until    the   iinilecnlcs   in 

"hitiiin    have    licciniie   ilissucialiil.    when    each    inn    carries   a    i-haiife    nf 

■  I'liiisiii'   si'jii    I  see    patre    llli.    whcriiis    in    the   case    nf    cnllniils.    each    enl 

'■■111  pMilicle  iisiially  carries  a  cliarce  which  is  ah\a>s  nf  nnc  sinn,  cither 

(•■■siiivc  nr  ncnatixe.     ('nllniils   may   tlicrcfnre   he   u'tniipcil   intii   pnsitixc 

.iiiil   iie'.'Mti\c,  accnnliii!;  tn  the  cliari.'cs  which   they   carry,  ami   there   is 

.1  iliml  '/iiiiip  in  which  the  ehar!;e  nia\   he  either  pusitixe  nr  iicmIim'  ac 

ni.liiiir  In  the  nature  nf  the  ilispersinii  iiieiliiim. 

.\  inljniil  lint  ciirryiii'j  a  chari;c  tn  lic;;iii  with  can  he  eailscil  tn  assiiine 
•ui-  li\    the  actinii  nf  elect  Inly  tes,  fur  the  electrical   prnpcrtics  nf  cnHniils, 
-  \m1I  as  tlinse  nf  inert    pnwilers  suspcinlcil   ill    w.'itcr.   ale   reailily   in 
'uiiHiil    hy    tile   cli.irires   present    in    the   inns   nf   the   ilispcisinii    liicilinMi. 
Till'   II-  ami  (Ur   inns  are  espcciall\    lialilc   tn   c\erl    this   intliieiicc.      The 
iriiejcs  nf  inert    pnwilers   in   siispeiisiniis  ikanlin,   sulphur,   etc.i   carr>- 
!"■    ti\c  chaiL'e  when   the  water   in   which    they  arc   siispemlcil    is  aciili 
■I.  ami  a  nc<r:iti\c  charu'c  when  it   is  iiiailc  alkaline.     In  u'cncral.  it   iiiav 
^.ml  that  siispcMsiniis  nf  must   pnvvilcrs  ami  nf  iiisnluhlc  nr'.'anic  aciiN 
.Miter  I  c.  I.'.,  charcnal.  ccllnlnsc,   Uanlin.  c,is'inn'_mi,  m.islie,   free  ,ieii| 
"ii'.'n  leil.  etc.)  arc  elect rn-iic!.'ati\c.     (  M"  true  cnllniils  ferric  hyi|rn\ 
leri'iiiii  ilialysatuni  I    ami   scriini   •;lnliiiliii   are   pisiiivi'   in   aciil  sniii 
-:  aisi'iiiniis  siilphiilc   ami   scrum    •^Inliiilin    are    ncii.ilixe    in    alkaline 
■  !inii,  ami  scniiii  <;lnliiiliii  in  neutral  snliitinns  has  no  cliari,'c. 


MICROCOPY    RESOLUTION    TEST   CHART 

ANSI  and  ISO  TEST  CHART  No    7 


1.0 


I.I 


1.25 


|-ia 


2.2 


::  lis    IIIIIM 


1.4 


1.8 


1.6 


^     APPLIED  IIVMCZE     Inc 


1"6)    4Bi      0300       i'hoi 


r»« 


I'ln  Midi  IIIMIIM.    IWSIS   (IK    I'llVMIll.dlillAl.    i'i{(i(i:ssi:s 


'I'll  ,iM-c'i't;iiii  the  ii;iluic  ol'  the  clijiri.'-i'  Viirious  iiicIIkkIs  iiiiiy  Ik-  ciii- 
|)iip\cil.  Ill'  wliicli  till'  liilliiu  iiii:  jiic  iiiipiirlMiit : 

1.  Till'  iiiflliuil  of  I  li  liriijiliiin  SIS.  Tilt'  iMillnid  snlutiiiii  is  plncfil  in  ii 
I'-tillic.  liii'li  side  .iT  wliicli  (■iirrics  ;i  pliitlMiiiii  cU'rt imlc  ilippiiiu'  into  llic 
siilulinn.  Allrr  ii  Nllnliy  rmit  iliUiills  I'li'rtric  clllli'llt  llils  lit'cii  ;illii«cil 
til  jiiiss  I'm-  sniiic  tiiiic  tliiiiiiu'li  tlic  MiliitiiiM,  it  will  lit-  t'liiiiiil  that  tlu' 
fdlhiiil  (•(illci'ts  ill  till'  MiKiilc  (  « lii'ic  till'  ciirfciit  I'litcrsi  wlifii  it  is  :i 
iicL'ati\c  I'dlldjil  isiiicc  uiiliki'  clfctiji-  cliarui's  attrai-t  cai'li  ntlicr),  ami 
at  till'  i-alliipili'  hIii'ii  jl  is  pnsitivc.  In  tlii'  cas"  iif  culnri'il  snliitiims,  tin- 
Miiijiaii an  ln'  iiaililv  seen,  lint  utlii'iwisc  it  may  lie  in'ccssary  tii  ana- 

\y/.f    lllr    Milliliiill    at     lllr    tun    pull's. 


,1,      ''-        .'  'l     'I"'       .l.i.'l..-.-      nl       ,,„|..„U,  S,,,|..      ,,1       |,1„,       |,,-,|„.,-.      ,,l„r       l.,i,l«      M1S..M.I.-.I      «,.l, 

"''"''    •'"'.'"'    '<-■    '<'    ill.      -ini-.    I", I     li.r.iu.l    a    >l,ari.    li,„     .„     ,1,  ,„ar>  all,.,,    at    lllr     |..w.r    .11.1     „„ 
■'"""'        ■'     l""l"I.lli"ii  lh,,s,     ..„    t|„     K,,    |..„„i     „,.,,.    n.^.uiv,.    ..,l|,,i.|,.        ||'n..ii     W.    IMtt,.l,|    ■ 

•_'.   Till'  nictliiiil  III'  ciiiillliirii  iniiihisis.     j-'in-  iliis  iini'iiiisi'  a  Iniifr  strip  ui 
liltir  papiT  is  aiiaiivM',1   \  tiiii'iilly  iimt  tlif  s.iliilion,  with   its  liiwcr  cinl 

•''PI'Iiil:  iiitci  it       In   ili,.  ,',is,.  ,,f  ni'-ativt Ilniils  the  ciilliiitl,  as  wt'll   a- 

till'  i|i>piiM.ii limn,  lisis  nnil'iiiiiil\ tlu'  strip  of  paper  (it  may  lir 

111   a    liiiu'lit    111'  20   cm   .  :    «  In'ifiis   with    pnsitivi'   I'lilhiiils    the   ilispi'isimi 

"""'''I'"   •'! '•!-''-.   Ill''  I'l'lluiil    itsi'lr  ilnimr  m ly   In  a    M'ly   sliM|,t    ex 

tfiit.   Imt    lii'i-iimiiiLr  s,,  lii'jhly  .-m nlratfil  at    tlif   int.'rl'ai-f  lii'l\M',.n   tiir 

siiliitiiin  anil  tlir  papi'f  tli;il  it  ciiii.rnlatfs  mi  tlii'  t'lnl  nl'  the  strip  ,,f  pjipm, 
wlirri'  it    ruiiiis  a  sharp  line  ut'  iliimin-atinn  (  I-'Il'.    1.")). 

:i.   Tlif   iiii'IIhmI   lit'  iiiiiliiiil  i>ri(iiiH,ill„ii   111'  fiilliiids.      When  a    ptisilivr 


CDI.I.OIDS 


/ 


1 


;iiiil  II  iM'Kiitivf  cDllnid  iiic  tnixcd  in  such  inoportiniis  that  llic  <'lccti'it' 
i-hMi'jjt's  arc  nciit  lali/cd,  |)i'cci|iitaliiiii  usually  occurs.  Wlicu  it  docs  so, 
uc  can  tell  the  iiatufc  of  tlic  cici-tric  cjiai'jic  of  an  unknown  colloid  hy 
il>  licha\ior  when  a  colloid  of  known  clcctiic  siirn  is  added  to  it.  For 
c\ain|ile,  if  ferric  hydroxide  (positive'  causes  a  piccipitate  to  foiin 
when  it  is  addi'd  to  an  uiduiow  n  co||oi(lal  solution,  the  clcctiic  charirc 
uf  the  latter  niusl  lie  nc"jali\e:  if  it  docs  not  precipitate  with  iVr-ric 
iiydioxiile.  hut  docs  so  with  aisenious  sulphide  ( iieirative  i,  it  must  he 
|iiisilive. 

5.  Brownian  Movement.  Like  the  paiiieles  In  lini>  niechanical  suspen- 
sions, those  of  colloidal  solutions,  espeeiall.v  when  exauLined  ultra- 
iiiicioscopicall.x .  exhiliit  the  so  ealleil  Hrownian  niovcinents,  which  have 
lieen  dcscrilied  as  "daiiciiii;,  hopi>in^'  and  ski|ipin'_'."  These  uiovcuicnts 
nciur  in  straitrl.t  lines,  which  are  siiddcnlv  (duinired  in  direction  and 
,ii-e  ijuitc  independi'lit  of  exteiiial  sources  of  enerjiy,  sucli  as  chau<rc  ill 
liiiiperatiire  (altlioi|M||  ijicy  hccoine  <|uicker  as  the  teinpcratiire  of  tile 
solution  is  raised*,  earth  vihrations,  chemical  (dianu'es,  or  the  electric 
c''Mri.'e  (d'  the  <'o||oid.  The  iiiov  ('iiicnts  liecome  iiioi-e  lapiil  the  smaller  the 
particles,  and  they  lieeome  sliie<;ish  as  the  vis<'osity  of  the  solution  in- 
iieascs.  .\d<lition  of  electrolytes  deeieases  the  movfiiicnt  liy  callsiiiu;  the 
p.irticlcs  to  clump  toirethcr.  The  density  and  viscosity  of  the  disper- 
sion medium,  tiie  electric  cliaru'e  of  the  dispersoid  and  the  presi'uce  of 
llnnv  iiiaii  movements,  are  the  forces  whicli  operate  toe:cther  to  prevent 
SI  cliiiieiitation  of  the  ))articles  in  a  colloidal  solution. 

6.  Osmotic  Pressure.  As  one  of  the  disiin^iiishiii<;  properties  of  col- 
hiids  uc  have  seen  that  their  dilTusiliility,  as  into  <relatin  or  aiiar  ,jel- 
!h's,  is  extremely  slow  when  coni|iarcd  with  that  of  a  mole<Milar  solution. 
Tills  does  not  iiieaii.  however,  that  colloids  are  possessed  of  no  poxv  er  of 

liffilsihility   if  left   lolii;  enoULdi.      Indeed   thi'  existeiiee  of  the   ilrovv  iiiail 

iii\tiiiciit    indii-atcs    that    siudi    ditTiisioii    i\[hxI    occur,    and    theiefon'    it 

-!  iiuld   he  possilile,   liy   the   application   of  the   same   piiiicipli's   as   those 

Mch  i;o\ern  molecular  solutions  le.  ir..  hy  usiny:  a  scmiperiiicalile  mciii- 

iiiei,  to  measure  the  osmotic  ))rcssu!-e. 

M.iny  studies  of  the  osmotic  properties  of  colloidal  solutions  have  heeii 
'■   'I'llaken,   especially    hy    those    who   are    iiitcii'sted    in    the    |)nssiliilit.v 

'      '  till Iloids  of  lilood  scriini  (serum  alluiniin  and  ylohuliii  )   ma.v  cre- 

.111  osmotic  pressure.      If  this  should   prove   to  lie  the  ease,   it    would 

•ei-i'Ksary     I'or    tlle    osmotic    pressure    |o    lie    overcome    hy     iiieclianieal 

-siiie  siirli  as  that  supplied  li,\    the  heart    (i.e.,  the  lilood   prcssiirei    in 

;iiioiis  pliysio|o<;ic  pidecsscs  of  liltraiiili  and  ditl'usion   lakilii;  place 

■  ■:\:  cell  uieuihranes  (as  in  tlie  formatioii  of  urine  in  the  ludney". 

'  f  iiicasiiriMc:  the  osniotie    pressure  of  colloids,   osinonieters   similar 


j..^i 


58 


rilYSI(UCIli;.MI(AI,    IIASIS   OF    rHYSKM.IHilCAI.    l'K()<'i;SSI'.S 


ti)  those  iiliTiidy  tlcscrihnl  I  li;if.'f  4)  cilll  lie  ciiililiiyiil.  Most  of  llif 
rcM-cnt  work  lias  liccii  done  citluT  with  colliHlioii  sacs,  or  with  iiiiK'n/oil 
(•la\-  cups  iiii])rc<;iiat('tl  \xith  sonic  ccl,  such  as  silica  i  r  jjclatin.  When 
siicli  ail  osinoiiictcr,  liMcil  with  some  colloidal  solution  (like  a  solution  of 
pure  ailiumiii'i  and  provided  with  a  vei'tical  <,'lass  tube,  is  ])laced  in  an 
outer  vessel  eoiitainintr  water,  the  lliiid  will  lie  seen  to  I'isc  in  tlie  ver- 
tical tulx',  the  height  to  which  it  rises  licinj;  ))ro|iortional  to  the  ostiiotic 
pressure. 

r>nt  tlie  observed  pressure  does  not  necessarily  iiive  us  the  osmotic 
Itressiire  id"  the  jiiirc  colloid.  I'oi-  to  tiiis,  even  wiien  hijrhly  imrilicd,  there 
is  almost  crtaiii  to  he  attached  a  euiisidcralile  aiiioiiiit  of  iiiorfianii- 
salt,  which  may  he  res|»oiisilile  i'nr  the  osmosis.  II  lias  indeed  lieeii 
maintained  by  some  oliservers  that  electrolytes  form  an  intesxral  part 
of  certain  colloids.  beiii<r  bound  to  them  ])erhai)s  by  ailsorptioii  (sec 
pai.'e  (i.">),  and  thai  lliey  are  es,seiilial  to  the  maiiiteiiancc  of  the  enlloid.-d 
state.  In  any  casi-,  since  electrolytes  are  .•ilw.'iys  present,  the  osmotic 
|iressurc  of  the  imre  colloid  ciiii  be  iiK'asured  only  when  means  are 
taken  to  discount  their  inlluencc.  Scver'al  devices  have  been  used,  of 
which  the  follow  iiij;  may  lie  mentioned: 

1.  .\dditiiiii  to  the  tluid  outside  the  osmometer  <d"  a  |>erceiita<;e  of 
salt  e<|iial  to  tluit  found  by  cheiiiie;il  analysis  to  be  present  in  the  col- 
loid.    (This  method   is  uiiti'ustwortlix .  ^ 

2.  The  use  of  a  limited  <|u;iiitity  of  tluid  on  the  outside  of  the  osmom 
eter  so  that  e(|uality  of  salini'  content  soon  iiecomcs  established,  b\ 
difTusion.  in  the  tliiids  on  the  two  sides  of  tlie  iiiciiibraiie. 

'■'t.  The  use  i>\'  a  membrane  which  is  peimeable  to  electrolytes  but 
not   to  colloids. 

Kveli  when  the  jrreatest  caie  is  taken  in  its  measurement,  the  osmotii- 
pressure  of  a    iriveii   colloid   lijis  been    fipiind    to   vary   considerably   not 
only  accordiii!;  to  thi'  iiu'tliod  used  in  its  preiiaration.  but  also  ai'coid 
in<r  to  the  amount   of  mechanical   .•lu'itation   (shaking',  stirrintr,  etc.)    t^i 
which    the    colloid    solution    has   been    subjected.      Ke};ar<linsr   the    inllu 
once  (d'  the  method  of  prcpjiration,  if  was  t'oiiiid  in  one  series  of  experi 
lueiits   that    albumin    that    had    been    repeatedly    washed    (but    still    con 
taineil    considerable   aslii    triive    no    osmotic    jircssure,    whereas    anotln  ' 
pi'cparatiou   that   liiid   been   ])iiri(ii'd   by   crystjillization    iwice   (and   con 
taincd   much   less  ash*    liacl   a    luessure   of  M.oS   mm.    11^.      Accordiii<r   !• 
these    results   the   ash    content    of   the    eollni.l    is    not    fundamentally    ii 
sponsible    for   its  osmotii-   jiressure.      As   to   the    iidluence   u{   mechanic, 
agitation,    the    osmotic    iircssure   of   a    {iclatiii    solution    is    increased    i' 
shakitiir.  while  that  of  a  solution  of  eirir  albumin  is  decreased. 

The  property  upon  which  the  osmotic  presstire  depends  is  tindoiibtedl 


IDl.l.OIDS 


59 


llif  stiitc  111'  dispcisioii  ..I'  tlic  colldid  iiiirtii'lrs,  iiiul  (liitil  we  Uninv  all  of 
ilic  fiiclors  wliifh  iiiiiy  iiillin'iicc  this,  iiiciisiiiciiii'iits  of  osiiiniic  pifssiircs 

III'    Cllllll 

|ii(i|)('i'ty  (ills  sonic  |ili\  sio 


ids  ciin  sciiicciv  l)i'  of  vci-v  iimcli  \:\\w.     N'cvcitliclcss,  lliiit   this 
iosrii"  hcaiiiii.'  is  dear'  from  the  <'tTc('t  which  col- 


iiavc  ill    rcstoriiii.'  tli<'  lilood   iircssiirc  after   licinori'liai;c    (  pairc    141). 


{•'ui-thcr  cxidciicc   that    the   osmotic   jircssurc   o 


f  colloids   lias   not   the 


siirnilicance  llial  it  has  in  the  case  of  molecular  solutions  is  furnished  liv 
the  fact  that  the  osmotic  i)rcssure  is  only  ap|)i'oxiniatcly  i>roi)()fti<inill 
to  the  concent  lation  of  the  sohition;  it  may  either  increase  or  decrease 
relatively  to  the  slrenjjth  of  the  solution.  Temperature  also  has  (piite 
a  dilTerciit  inllnence  on  the  osmotic  pressure  of  colloids  from  that  wliicli 
it  has  on  the  osmotic  pressure  of  molecular  solutions,  and  it  fropu'iitly 
lias  an  indiience  which  persists  after  the  solution  is  hroiinlit  hack  to  its 
iiiitrinal  level. 

The  inllnence  of  aiided  sul. stances  on  the  osmotic  jiressure  of  colloidal 
siilulioiis  is  of  coiisi(|ci-;l)le  interesl  to  the  liioloeist,  for.  whereas  in  the 
iMse  of  •  ilecular  -^i  liitii  ns  this  is  purely  additive,  in  the  case  of  col- 
Inids  the  add(d  snhstanee  may  at  one  time  cause  the  osmetic  |)ressiire  to 
increase,  at  another,  to  decrease.  It  has  hecM  found  that  the  osmotic 
piTssiire  of  frelatiil  solutions  at  first  decreases,  then  rapidly  increases  as 
ihc  ll-ioii  ciniceiitiation  is  raised.  The  additii  ii  of  alkali  increases  the 
•Miiotic  ])ressure  until  a  maximum  is  reached,  licvoml  which  it  hciriiis  to 
I'.ijj.  I'xith  acids  and  alkalies  h'ssen  the  osmotic  pressure  of  eir!.'  alliu- 
iiiiii.  KIcctrolytes  always  decrease  the  osmotic  lu-essuie  <d'  trclatin  and 
illmmin  solutions,  and  the  deirree  to  which  they  exert  this  inlluencc 
ill  pi'iids  on  the  nature  of  the  cation  and  anion  composini;  the  elcctrolyt*-. 
Ill    the   order   of   their   de|)ressiiij;    intluence    the    ((iliniis   aiTanH:e    thcm- 

lieav.v   metals  >  alkaline  earths  >  alkalies; 
■  iiiil  the  fniii)iis: 

SO,  >  V\  >  NO,  >  Hi-  >  1  >  ("NS. 

ill'  intluence  of  a  ^'ivcn  electrolyte  varies  extraordinarily  with  the  reac- 
iii  of  the  colloid,  a  fact  which  must  he  carefully  re^rarded  in  all  work 
;i  tliis  field. 


rll.MTKI.'  VIII 
(OLLulDS  (('..iitM) 

SUSPENSOIDS  AND  EMULSOIDS 

Accdrrliiiii    til   \\  lictlici'   CMlluiils    t'ciriri    M.liiti(p|is   tjuit    .-lie    i c   u\-    |,.ss 

viM-i.l  tli;iii  the  Mis|H'iiNi.ifi  iiiiMlliirn.  tlii'x  .iir  ijiviilcd  into  i  niiilsniils  hikI 
.■<iisii,  ii.saiils.  K\,iiM|.li'N  „(  \hv  r.iiiiicr  clii^s  ;nc  sili.Mtcs  iiikI  u'cliitiii,  iiimI 
of  thr  l.ittfi-.  .|i,il.\/('<l  iron  iiikI  iiisi-riimis  Milpliidc.  Tlif  ruling  iiij:  cli.ir 
iicli'ristics  lire  iiscil   to  distiiiu'iiisli   Ix'lwccii   siispcnsnids  iirid   ciimlsiiids: 

I.  Mi'.iMiiiiii:  till'  time  it  tjikcs.  ;it  ;i  st,iMd;iii|  t.'iii|pcriitiiir,  t'nr  a  ^iviii 
viiiiiiiif  «(  tiic  lliiid  to  How  out  of  a  staiiilard  pipctti-  I  10  (•(•.)  slious  tlic 
risnislhi  to  he,  roiiuhl.v,  invi-rscly  iM-opoiiiuiiat  to  tlir  tiiin'  of  oiit(l<i\\.     In 

'' •1'^''  "'"  MisptiiNoids  till'   vis<-osit.v    is   no  dilTcn-nt    ftoni   tliat    of  tlo- 

dispirsion  niriiiuiM  aloiir,  and  does  not  vary  niiK-li  when  the  solution  is 
i-oolcd.  Tln'  viscosity  ot'  cinulsoids  even  in  vi'i-y  dilute  solutions  is,  on 
the  otlier  hand,  eonsideialily  u'reater  than  that  of  the  disi)ersion  ineijiurn 

itself,  and  it  lieeornes  !.'reati>    iiiei-eased  hy  ilinir. 

'_'.  Suspensoids  are  niueh  nioi<'  rea<lily  eoa!.'ulafed  liy  tlu'  addition  of 
eleetrolytes  than  eniuls  ids.  This  is  ))artieularly  true  when  water  is 
the  dispersion  medium  f  so-called  hydrosolsi,  ami  when  eleetrolytes  ha\ 
inv' a  polyvalent  ion  (such  as  Al  or  MfS.)  are  enii>loye(l.  Thus,  practieall.v 
all  suspensoids  are  eoaj^nlited  in  the  presence  of  1  per  cent  id"  alum, 
which  has  no  inlluence  on  cmidsojd,.  We  shall  return  to  this  phase  ol 
our  suhjecl    l;itel-  on. 

The  division  of  colloids  into  emnlsoids  and  suspensoids  is  more  or  less 
arhitrary,  since  one  class  imiy  lie  ehany:ed  into  the  other,  tlw  determinin',: 
factor  lieiuL'  the  water  content  of  the  dispersoid.  The  water  content  oi 
susi)ensoids  is  low  I  lyophohci,  while  that  of  emulsoids  is  hi-rh.  I'.n 
ehanL'in.r  the  idalivc  amounts  of  w.-iter  and  solid  of  which  a  colloida! 
.solution  is  compiised.  the  nature  of  the  dispersoid  may  he  chanireil.  I' 
the  water  is  diminishrd.  the  dispersoid  lieliavcs  as  ;i  suspensoid  and  he 
comes  r.-ailil\  pncipil.itcd.  The  practical  importance  of  this  fad  i^ 
that  il  explains  tin  sulliiui  „iit  „f  in-nhiiis  a  process  extcUsivelv  usc.i 
in  their  separation,  (trdinarily  these  liehave  as  emulsoids,  hut  the  add) 
tion  of  salt  raises  ihe  nsmolic  pressure  of  the  dispersion  iiu'dinni,  aii.i 
thus  attracts  water  from  the  dispersoids,  with  the  result  that  they  com. 

I  ill 


CI  11, 1. 1 II I >s 


61 


ici  licliaM'  !is  siispciisoids,  iiiiil  iiri'  iiiTuiiliin.'ly   pici'ipil.'itcd   \>\    \\w  clcc- 
licilx  Ics. 

AnnllH'r  |iiii|ii'ily  nl'  I'lmilsniils  nl'  liii>lii<;iciil  im|Mirliiiicr  is  l!ic  jiro- 
Iirlioii  whii-li  tli('>  cjiii  iilTiinl  imiiiiisl  llic  pi-ccipilMliiiy;  iiilliii'iicc  id' 
rli'fti'iil.x  ti's  1)11  siispi-iisiiiils.  II'  :i  i-iilldiiliil  siilutimi  nl'  unlil  is  niisi'il  with 
a  liMi'i'  III'  jidiilin,  llic  Milisci|iirlit  iiiliiit  inn  nl'  suits  x\ill  lie  fnuilil  to 
I'liiilurt'  no  prrripitatiiiM.     Tlif  rxplaiiiitimi  nl'  tliis  is  tluit   tlic  I'linilsniil 

III iiics  (listriliiitril  as  a  (iliii  mi  tlir  siispi'iisniil  partii'Irs,  tiiiis  iiiartii-iilly 

■•iiiiv crtiii!;  tlii'iii  into  I'liiiiisoiils. 

Gelatinization 

(•ill-  III'  till'  lirsi  Iviiiiwii  ]iniiiiMtirs  III'  I'liiiilsiiiils  is  tliat  of  u'l'latiiii/.a- 
liiiii.  wliirli  lias  an  iiiti'ii'stiii!.'  lii'ariim;  mi  iiiaiix'  pi-ulili'iiis  <>(  liiulmry. 
AfliT  till'  u'l'l  lias  set,  an  I'limiiimis  pii'ssmi'  is  ri'ipiiii'il  tn  sipii'i'/.i'  mit 
,iiiy  wati'i'  I'l'iiiii  it,  iiuliratint:  tliat  tlii'  watrr  iin  Imiv'i'i'  riniiis  tlio  ciiii- 
iiiiiimis  ]iliasi'  liiit  must  In-  I'lirliisrd  in  vi'sii-lcs  t'liniii'il  nl'  niiirc  siiliil 
iiiatt'iial. 


As  a  ;.'rlatili  snlutinii  riinis,  tlir  fxr]  at  lirst  I'liinis  a  |inl;ni/i'il  riilli'  nl' 
:lit.  lint  till'  vi'i'v  liiif  partii'Irs  wiiidi  an'  I'i'spnnsililt'  I'm'  this  I'lTi'i-t 
■11   iiici'i'ast'  in  iiiiiiilii'r  and  s'y/.v  sn  that   thi'y   nlistriii't  mii'  aiiotlu'r  in 

■ir   iirnunian    iiinvi'iiit'iits  anil   ailhi'ii',    yiviii'^   an    a]i|)('aiaMri'   nf   fiiii' 

t  lil<i'  thi'oails  tlirmii;hnut  tin'  snliitimi.  A  smt  nf  iinpiTvimis  spnii^i' 
''il<  nf  tlio  liinrt'  snlid   pliasi'  is  thi'iffni'i'   fniiiii'd,  thi'  iiinri'  llnid   jiliasi' 

iii'j  iiH'lnsi'il  in  till'  inrsht's. 

1 1'.  as  in  till'  ai'i'iiiiiiianyiiiK  ilia>;taiii,  tin-  dispi'isimi   nn'diiiiii   is  icpii'- 

■ 'I'd  liy  white  and   tin'  dispi'isnid   in   lilai'k,   tlir   ri'latinnsliip   liotwi'i'ii 

'  ■■•>  in  a  siispcns.iid  is  as  in  .1,  and  thai  in  a  fill  as  in  /.'.     Tn  i\|iiiss 

.    I  I'  till'  dispi'i'simi  iiii'diniii  in  /{,  it   will  ri'iiniri'  a  pirssiiri'  siillii'iriit  tn 


(IJ 


I'llVslriirlllMlc'Vf.    iiAsIs   oK   I'll  Vslul.iiiilr  \l.    I'ldicilssi'S 


iMllv    llic    IMKIi'    ttlliil    |i|i;isi'    111    lirlifli.ilc    llir    iiiiiic    siiliil.       It'    the    tfcl    is 

liTjiti'cl  with  ic.-iiri'iil.s  liki'  ri>i'iii.iMi'li\.lf,  the  lii|iiji|  i-jin  lie  ifiiilil.%   in'cssfil 
mil.     'I'liiy  iK'riirs  (liiiiii"^  tiv.itiiin  I'nr  )iist(i|ii;ric;il  |iiir|).isfs. 

Imbibition 

Clciscix      ri'ljili'd     tn     irrl     I'lillllilt  lull     is     the     |Uihtss     ii|"    iliiliiliit  inn       111!' 

|ii(\M'r  r.r  tiikiiisj  ii|i  liiisic  <|ii,nilitirs  of  \\:\\i'v  willioiil  ii''tiiiilly  roniiiii!.' 
Ii>|uii|  MiliiliiiMs.  Mcsidcs  i:cl;iliM  tlic  dlicil  tissues  ipf  pliilits  iiliil  iiliilliills 
('\liiliit  tlic  iiliriinini'iKiii,  iiiiil  it  is  iiiiilniiliti'illv  lit'  iiiipiM-taiict'  in  iiiiin.v 
plivsiiiliitric  piiicfssrs  siirli  iis  irinutli  ;ilicl  tlic  pMssnu'c  "f  Wfitci'  ilitn 
iM'lls.  etc.  'I'hi'  lii;it('li;ils  lilTsclit  ils  VJicilcili's  in  pl.illt  i-clls  .-ittiact  \\i\\v<- 
t'rniii  till'  I'lu  ii(iniii<iit  I'f  till'  fill  liv  iiiiliiliitiiiii,  miii  thus  rxcif  i.ii  tin' 
'•I'll  Mall  a  iircssnii'  wjiii-l',  artintr  alniifr  witli  llir  nsiiiotic  |)ri'ssiiii'. 
iiiaintaiiis  llu'  tiiiirui'  nf  tin'  I'vil.  'I'lic  initial  triowtii  of  piillcn  is  alsn 
lii'pi'iiili'iit  upon  iiiiliiliitiiiii,  ami  inipnitant  ol)s('r\  atinns  nii  tliis  jn'oi'i'ss 
iiinliT  \ai\in<:  ronilitinhs,  air  liki'ly  to   I'mnisli  us  witli  nscfiil   infoiiiia- 

li"i iii'i'inin'r  tin'  siLriiilirain-i'  ul'  iiiiliiiiitinn  in  I'linni'rliiiii  willi  (fi-nwtli 

III'  fi'lls  in  irriiiial. 

Uy  iiii'asiiriii".'  tlic  rati'  «(  inrri'aM'  in   li-n^rtli  of  Imiv'.   iiarrnw  strips  nl 
U'l'Iatiii   plai'i'il   ill   I'l-tii  ijisln's  mntainin'r  soliiliuns  of  vaiyiii>r  i-innposi 

tiiiii,  the   fai'tiiis  tliat    iiitliu'iii-i'  tlii'  iiiiliiliitinii  jiii ss  i-aii   lie  ipiaiifita 

livi'ly  invi'stiiratcil.  Wiirkini;  in  iliis  way,  I''.  II.  Unyil'"  has  I'lniinl  that 
I'nr  all  ai'iijs  tlii'i-i'  is  a  i-i'rtain  I'mi'-rntratinn  (almnt  X  :!-2(l  II.SO,)  wliirli 
iiiiliii'i's  a  iiiaxiiiiiiiii  rati'  nl'  svvi'Hin>r,  ami  aimtlii'i',  iiiiii-h  wi'aUi'i 
(  .\  •JSdtl  II.S(»,i.  in  which  tlir  rati'  of  s\vi'llin>r  is  I'Vi'ii  Irss  than   in  jniii' 

"ati'r.     In  lii<;hi'i iii-i'iitratiniis  nf  ai-iil  than  N   :{-2(l.  tlu'  <ri'latiii  at   liist 

sMi'lls  vi'iy  i|uii'kly.   lint   the  rate  sinws  nfl"  sn  tiiat    it  smpii  I'lniii's  tn  I" 
less   thai,    that    with    inli'iini'iliati'   rmici'iitratiniis.      Hcsiarilinir    alkaiii's. 
at  hi-ih  I'niii'cMtratiniis  the  I'lTcrt   is  similar  In  that  nf  aciils.     Salts  ahun' 
si'i'iii  tn  ri'pri'ss  till'  swcllinir  lii'lnw  that  ni'  wati-r.     It  slnmlil  In'  pniiiti'<l 
nut  that  the  i-nnrcntratiniis  nf  ai'iil  ami  alkali  in  tin-  almvi'  nhscrvatinii  ■ 
art'  iiiui-h  L'li-atcr  than  tlmsi-  that  i-nuhl  nci-ur  in  tin"  animal  linily.     Tli' 
I'xpiTiiiii'iits  ii'i-all  till'  attempts  maile  smiie  years  afin  hy  Martin   l'"isehei 
tn   explain    eih'iiia    as   ilue    tn   exeessive    imliiliit  inli    nl'   water   liy   the    pin 
teiiis   nf   the    tissues    lieeai'se    nf    iliereaseil    aeiility    nf    tile    lilnnil    ailil    li- 
siie    tluiils.      That    iiiiliiliitinii    miu-ht    pnssihly    ]>lay    snme    rnle    in    sinl 
prneesses  is  nnt  ilenieil,  hut    l''i>eher  ilisre-ranls  entirely  the  iinw  well-i'stal' 
lisheil   faets  that  li\  ilrnneii  imi  eniiceiitratinn  is  nne  nf  the  liinst  ennstan 
pinpeities  nf  the  lilnnil,  that  \  ery  Inu   eniii'i'iitiatiniis  nf  aeiil  may  iliiiiin 
ish    rather  than    inerease   iiiihihitinii,   ami   that    it    is  manifested   nidy    i 
the  ahsenee  nf  innruranie  salts.*     Mnrenver.  the  ttiiid  in  ede  iiu  can  eft' 

•I>,l,,,„,„,-,t„.„.   ,,t    th,     l,x,!,.^.,  n  !..„   ,..,!..  1,1. .Hi f    ll„    1,1 1    ,,,,„||v    |, 1,1,1, .li,,l    f,,„„    |.„,h, 

l;,!„,i;,t,.ry   ,!,,   ,i,.t    i,,-i.,i,    ,  ,,nr,.l!  m,  tv 


l'OI.I,lll|iS 


f.;j 


l.r  ilriiiiK'il  (ilT  \)\  liiillou  nci'illcs.  iiiiil  it  piisscs  l>y  jrravity  finm  <mc  jiait 
..I  the  liliiitil  1o  aiKillicr,  ncinici'  nl"  wliicli  in-occsscs  winild  bo  ixtssihlc 
ii  iiriliiliiliiiii  wcri-  the  cssfiitial  fat'tiir  cnMCfnifd.  If  I'iiiIIh'i-  cviilciici' 
iiHiiinst  tliis  tiy|)(illicsis  slmulil  lif  ilciiiaiiclnl.  it  iniiilit  lie  t'oinul  in  tlif 
iltiT  failiiii'  of  till"  tliciapfUli<'  imasnri's  alkali  ailniiiiistratiim  tliat 
III'   rrcoiiiiiU'll<lt'i|   Id  ciiiiiltat    llir  <'i|f|iia. 

Action  of  Electrolytes  on  Colloids  (aparl  frnm  tlifir  clTi'ct  i>ii  nsniotif 
IHi'ssiiii').  It  lias  liccii  statfil  aliiivc  that  the  cliar-.'*'  "Iiifli  a  ('"lliiidal 
|iiiiticlf  assiiini's  inav  Ix'  iicutrali/.cd  l)y  a  cliarsic  of  opposite  sij;ii  <'ar- 
I'ii'il  liy  an  ion  pr'cscnt  in  the  dispci'sion  nicdinni.  Tlic  ncntrali/.ation 
111'  till'  rit'ctrii"  cliar'iri'  canscs  coaiiidation  of  the  siispcnsoids  but  not  of 
till'  cninlsoids.  Of  the  positive  and  ne);ative  ions  into  wliieli  tlie  elo.'- 
iiiilytes  dissociate,  tlie  one  pi'odiieini;  tbe  i-oa<;nlation  is  tliat  wliieli  is 
ci|ipo>ite  in  sij;n  to  tlie  elei-tric  eliar^'e  of  tlie  eolloidal   pafliele. 

A  quantity  of  eleeti'olyte  wliieli  is  capable  of  produciiiy:  complete  |>re- 
ripilation  \\lieii  added  all  at  once  to  suspeieoiils  will  lie  inetTective  «  lieu 
Mililed  in  small  i|naiitities  at  a  time.  This  plieiiemeiiou,  wliieli  is  also 
kiiiiwii  to  be  eshibited  wlieii  toxins  and  antitoxins  are  mixed  lotrether,  is 
pi'iilialil.N  owiiiir  to  tlie  fact  tliat  precipitation  depends  on  ine(|uality  and 
iiirirular  distribution  of  electric  (diar<;es.  a  condition  wliieli  liccoiiics 
rNliililislied  wlieii  tlie  electrolyte  is  suddenly  added,  but  not  so  wlieu  it 
is  u'radiiaily  added.  The  iiaiti(des  ill  the  latter  case  be<'oiiie,  as  it  were, 
;Hiruiialed  to  the  electric  <'liai<;es  introtliu'cd  by  the  addition  id'  llie 
'■irctrol\  te. 

Proteins  as  Colloids.     The  most  ])romineiit  colloids  in  the  field  of  liio- 

i'Ih  iiiistry    are   the    jiroteins.      On    account    of   complexity    of   structure, 

!iii\\('\  cr,   certain    factors    intervene    which    render   the    iii\  cstij-'atioii    of 

:lirir  behiivior  vei'v  diilieult.     As  we  shall  see  later,   proteins  are  made 

'■i]>  of  combinations  of  amino  acids,  caidi  of  which  contains  basic  iNIT.! 

il  acid  irroups  (COOlI).     Tlie  various  amino  acids  are  lihked  loyietlier 

protein  by  the  COOll  of  (uie  unitiiur  with  the  N'll^  of  aiiotlier.  with 

'■    liiiiiinatiou  of  water     thus.  CO      Oil      11:     UN'     but  sume  \ll.^  and 

•     "dl   u'loups  are  left  uiieonibined.     Aceordinsr  to  the  i-clative  nuniber 

!    Iliesc    uncombiued    radicles,    the    imitein     (or    pol.vpeplid,    jtafre    (id!) 

'I  iNhibit   faintly  acid  or  basic  or  neutral  i>roperties.     With  acids,  f<u- 

.Miiple,  a  salt  will  be  formed  by  union  with  the  Nll.^  irroups,  wliiidi  will 

-M"-iate  int<i  the  anion  of  the  acid  and  a  larjre  or<ranie  cati<ui;  whereas 

Il  iiikalies  union   will  occur  with   the  COOIl   Lrn.n]).  and   the  salt   on 

i.itiiij;  will  form  a  small  cation  i>(  the  metal  of  the  salt  and  a  larire 

iiipiex   anion.      We   may   liierefore   obtain    the   lu-otcin    with    either   a 
-.live  or  a  lUfjative  electric  cliar<je  by  alterinii  the  eheiiiical  nature  of 


1 


til 


I'lIVslriM  III  Mir  M.    ii\sis  ok   I'll  Vslul.(HMr  \i.   I'IKxi  ssi  h 


till'   lliiiil    ill    whii-li    it    is  ilisMilv  I'll,   mi   lli.'il    tin'    ii'iirl  imi    tovMiiiis   otlii'i 
I'liJIiiiilN  Mini   tiivt  :ii'ii'>  rli'i'i  I  iil\  tiN  will   v.'ii'V. 

Olll'    I'l'Mtlll'l'   III'   |ll'l>tl'illN   III'   itll|lii|  tillll'l'    ill    this   I'lllllK'clioll    is    tllilt    klHlNMI 

lis  the  (Silt  li  ilrii-  iiiiiiil,  iit  wliirli  llir  |ii'iiti'iii  exists  with  ii  liiMxiiiilliii  iil 
I'li'ctrii'iillv  iii'iiti'iil  iiiiiU'i'iili's.  This  |Hiiiit  is  ifiii'lii'ii  liv  iiililiiii;  iti-id  In 
a  |>r<it('in  solution.  The  iicid  irpri'sscs  tlif  ilissiiciiitioii  of  tin-  j)i'oti'iii 
iii'tiii;;  iis  an  at'iii.  and  llii'i't'l'oi'c  diliiinislics  tlii'  niinilirr  of  I'rci'  |iydl'oi;ri> 
ions;  and  at  the  saini*  time  it  I'lUiiliiiu's  with  the  NIL  uroiips  and  im'IiIi'.'iI 
izcs  till'  liasic  idiarai'ti'iisticH.  Tlic  alteration  in  eleetrie  eliaine  tliiis  in 
tliieed  alters  the  \\  ater-alpsorliinn  powers  of  the  protein  and  tliefefori' 
all  ot'  the  priipi'ities  whieh  \\r  have  seen  to  lie  assoeiated  therewith 
I  pa'.'e  (i:;  1. 

SURFACE  TENSION 

I'.el'ore  We  consider  a  \er\  important  property  of  eolloii^;  known  a> 
atlsorptioii,  li\  means  ot'  whieli  the.\  are  aiiie  to  perform  liiali.\'  reactions 
that   do   not   eoiiforin    with   the   laws  of  mass  action,   it    will   lie   well    to 


I'tk'      ir        lM.tw":i'i)     I',     ilin^lra!)      siMl.ur     I.  iiviiin         Tin      liny-     ■'     aiiii     /'     iiitto-r     -iii.ti'  tiliii-i     iv 

uln.li    .1    %.M     t.iM     |...|.    .rl'    -ilk    I-    -ii-i \,.\        III    .1    il    I-    I.M.-r    lull    III    II.    »li.i<'    llii'    lillii  iii.|.>-i.i 

III    ill-      i""].    |i:,N    1„.ii    t,,.,l,.ii.    11     I-    ill., SMI    ml..    ;i    i  irt  It     1>\     llir    tiii-i t     III.     -iMp    Iiliii,  ll-'i^ti! 

K.nh"   1 


sav  a  lew  words  coiieeininLr  the  jihysieal  phenomenon  upon  which  tlii-- 
depends  namely,  surface  tension.  The  creation  of  this  force  is  die 
to  tli<'  fact  that,  whereas  the  molecules  within  a  liiplid  are  suli.jecled  t^ 
eipial  forces  of  attraction  on  .-ill  sides,  at  the  surface  these  forces  act  mi 
one  side  of  the  iiiolecules  only,  and  therefore  tend  to  jiuil  tlii'in  inwaiii- 

Tliis  causes  the  surface  to  |iull   it-elf  to«.'etlier  so  as  t ciipy  the  lea-' 

possilije    area,    and    it    is    this    force    which    constitutes    sui'face    tensimi 
The  siirl'ace  lieliaves  as  if  stretched.      There  are   various  simple  cniu'i  ■ 
ineiits  that   rcxcal  the  presence  of  surface  tension.      If  a  lilm  is  iiiaile  "' 
a   loo])  of  wire  liy  dippinir  it    in  soaj)  solution,  a   line  silk   tlircad  can   I  • 
lloated   in   the   lilm.  so  that    it    forms  a    loop  that    is  ipiitc   loose.      If  tl: 
portion  of  the  lilm  inside  the  loop  is  destro.ved  liy  toueliini;  it  with  lilt' 
jtaper.  the  lilm   will   lircak  in   the  loop,  which   will   now    lie  pulled  into 
circular  sliajic  liy  the  tension  of  the  film  around  it  ( l-'ii;.   17). 

For   till    tm  iisiirt  iiK  lit   of   niirfnit    li  iiaiiiii,    various   methods    are    iisr!. 


I'Ol.t.OIHS 


tJ.". 


'I'll.'  si/..'  til'  .liii|iN  (.r  lii|uiil   fiillinu   I'l'iiii  nil  ..lilif.'  is  .l.'|i.'ii<l.iil  »h  siir- 


lilO' 


Iciisi.m;  111.'  ImiKT  till'  .Imps,  (lif  iir.'iit.r  tlic  Mirf; I('Iim..ii 


111.'  riiiiiilit'r  (if  limps  nliliiiii.'.l  liv  iill.iw  iiiit  a  li.|iiiil  I"  .li"p  li'mi  n  stiiii.l 
,irtl  KriCK'c  in  a  (fivi-n  tiiiif  is  I'l.iiiil.ii.  \M'  liavr  a  iiiiasiiic  nf  liic  Mirtnff 


tciisiiiii.  Ai'c.iiiiit  iinist  tif  ciiiiisi'  alsii  lie  taUi'ii  of  llif  spn-ili.'  irraMty 
.if  till'  li<pii<l.  Tin-  iiistiiiiiii'iif  ns.'.i  f.ir  lliis  puip.isi'  is  .'aili'.!  a 
sl(il(i;iii)n>iiflir  (l-'itf.  IMI.  Amilli<r  iiiitlin.l  .Icpi'iuls  mi  tli.'  i'ai-l  tliat 
III.,  hfiiflit  to  wliicli  a  tlni.l  lis.s  in  a  I'apiilaiv  tnlx-  is  .l.p.  ii.l.tit  on 
siiifacc  Ifiision  (aii.l  iiiv.'is.'ly  .m  tlif  iliani.'t.'i-  ..f  the  .•apillaryi.  Tlif 
.lilT.T.'ii.'i'  in  till'  h.'it'lits  I.,  wlii.-li  \w>  li<ini.ls  lisc  in  .•apiilaiv  tiilics  of 

known   liorc   ptTinits   us  1 par.-  tli.'ir  siiifa.'f   1.nsi..ns,   an. I   if   this 

is  known   for  one  .if  the  s.ilnlioiis,  it   can   I"'  .i.lii  iiiiiii.l    for  tlic  otlicr. 

Besides  exis;intr  li.'tu.i'ii  li.iiii.l  ami  air.  surfa.'.'  t.'iisioii  alsi.  I'xists  at 

llie  int.'rfin-e  Itetween  Iw.i  iiiiiiiis'-ilil.'  li.pii.ls,  ami   al    tlial    lietweeii   sus- 


l,l,      IS        I'laul..'-     sl.il.iKiii.iiLil.  1.       ihc      -.iil.ii.      Iii-ii.ii     I-     i.r.iii..ih  'iiil     I"     lli.      niiiiili.  r     "f 
hn,.,    l,,rin,-,l    III    a    limii    Mi"  llu     uuiil  .mul. M    mlu^    an-     I..1     Ihiu     IlimmN.    an. I     lli.     sIuikIiI 

■  .»■    I.,t    hi I   an.l    i.l.i.r    m-...ii,    lUn.i>. 

iMMi.li'.l   solid   i)articies   and   li.piid.   as   in   .'.illoi.lal   soliili..iis.     Sin.-e,   as 
w  have  seen,  the  surface  area    i  iiit.'rfacc  i    is  en.iriiioiisly    increase.l   in 
hi'sc  solutions,  a  very  <in';\t  surfa.'.-  cncriry  is  present,  for  this  is  eiiiuil 
1  the  surface   tension   inulliplicd    liy   the   surface   area. 


ADSORPTION 

Till'   surface   tension   iM'tw.'en    /''/"/(/  mnl  nir   is   lowere.l    when   .ii'iraiiic 

Instances  are  diss.ilve.l  in  the  li<|uid,  liut   is  sliirhtly  raise.l  when  inor- 

-  mi.'  salts  are  dissoiv.'il.     The  iW'^H'v  of  ioweriiii,'  vari.'s  inarkeiily  ac- 

.idiiiLt  to  the  orjranie  suhstance  dissolveii,  li.'inn  very  proiniuiicd  with 

^  il  ■  Mills,  upon   whidi   fact  the  well-known   I  May)   test    for  th.    pr.'s.'ii.'e 

■  1'  Ml.'  ill  urine  is  liased.     I'.elwfen  //(/»/./  mnl  liiiiiid.  as  w.'ll    is  ...'tween 


1 


titi 


I'llSMi  III  III  Mil   \l     li\N|>    III-'    l'll\Mnl  iHili   \l.    riliH  I  ,ssi:s 


Mihil  mill  I'l/iiiil,  till'  siiiriMi'  ti'iiNicili  is  iihviuis  litinrtil  hi/  ilis.snlriiiii  siili 
xliniiis  III  llii  hiiiiiil,  N'liw,  111  till'  illlt'l  r.iiTs  lii'tWffil  siiliil  |ii|l'tii-li>s  jiliil 
liijiiiil  iliiir  iiiii'-t  Im'  ,1  liii-al  ni'i'iiiiiiiliitiiiM  iif  liTi'  Mifl'iiri-  I'lii'i'irv.  \\i.  .'Ii 
^<ill    III'    I'MUiil    1.1    till'    Mirl'jii'i'    tili>ii>li    liillltipliril    In     llic    Mii'l'iirc    (  iiiti'l' 

'•'I Illi'.l.       A    riiliNtiiiit    ti'lnlillrv    rxislx    t'lir   Mlrll    Iri'i-   nii'i'iry    to    III'    ill' 

I'l  •;iMil  ,iimI.  siiM-i'  ihssi.K  111  siilistiiiii'i's  liiuc  tliJM  fITi'ct,  tliry  will  Iii'imiiih' 
11  •ni'iilratril  ill   III!'  iiiti  rl'ari'.     'I'liin  is  kiiiiwii  lis  III!    iir'nii'ijiU    of   W'illittil 

tllhliX,    ilMll     It     is    111'     I  llllll.'lllli'Mtill     illlplll'tlllll'f    til     till'     Ililli-lll'llliHt,     I IIIIM' 

nil    it    ili|iiliiU    till'    |ilii'liii|iii'l|ii||    kliiiWII    IIS  Ili|Mil'|itiii||,    «liii-li    ill    till'    riisr 

111'  i'iiliiiiii;il  siil.itiiiiis  iiiiiy  tlii'ift'iiiT  hi'  (K'liiicil  IIS  till'  liinil  roni'i'iitni 
li'iii    111-    I'litnli'liMltiiiii    III'   ilisMilvril    slllistiiliiTs   111    file    iliti'i'riii-i<    li(>f  \\  fell 

llir  twii  |iluisi's.     Till'  ii lint  III'  siilistiiiii'r  I'liiii'i'iili'iili'il  lit   till'  iiitrrfai-'' 

'■iiii  I ali'iihili'il  liy  :i    I'liiimila   wliii'li  Ijiki's  iiitii  iici-niiiit   tin-  roiicciiti'ii 

liiiii  111'  III.'  i|isMil\ii|  siilislaniT,  till  ti'iii|)i  ratiiii',  am!  tlii'  siirt'ai'c  fi'iisimi 
at  till'  iiiti'i'I'aii'  I  till'  (lililis  I'lii'iiiiila  i.  AltiT  ali»iii'|itii>ii  lias  ni'i'iiri'i'il,  vai'i 
mis  I'l'ai'tiiiiis  iif  a  iliiiiiiial.  iln-triral  nf  |iiin'ly  |i|iysiral  iialiiri'  i  r. '_'..  ilit' 
riisiiiii  '  ,iia>  I'lilliiw  at  a  rati'  wliii-li  i|i'|ii'iii|s  mi  tin'  aiiiiniiit  nf  t' r 
rmiiji'iisal  imi. 

Everyday  Reactions  Which  Depend  on  Adsorption 

1.  DiTiilmi/atiiiii  111'  lli|iiiils  liy  i-liari'i>al.  Tliat  lin  clii'liiii-al  rrai'tioil  iii' 
I'liis  in  siii''i  a  ras-  is  rrailily  slmwii  liy  tin-  i-asi-  witli  u  liii-li  the  piu'iiii-ni 
ran  111'  iNliai-I.il  riimi  tlir  rliarrnjil. 

■_'  Ai|siii|iiiiiii  111'  "asrs  liy  siH'li  suliils  as  cliaiciial  iinil  s))m,-y  plaliiiiliii 
In  lli.sc  iMs.'s  ili-i'.'  iiiiisl  111'  liiTai  i-miili'iisatiun,  i  \i'ii  a  lii|ili't'artiiiii  nl'  tli. 
uas.  iliiriii'j-  w  liii'li  In-al  must  ln'  i'\  nlv  I'll,  \i\  ailsnrliiiiir  nxyt'i'ii  :  ml  liyili  n 
vii'ii,  spiiim.N  plaliiiiiiii  i-aiisi's  tlii'si'  L-asi's  tn  i-niiiliiiu'  anil  I'miii  wali-i'.  'I'lii 
liriiHii'liiliin  111'  liliiiiij  iiia.\   lake  up  usyji'ii  liy  a  similar  ihhim'ss.. 

:i.    I''iii'iiialiiin  111'  sujiij  Mii-laii'  Minis  mi  sulutimis  nf  piiiti'in,  sapmiiii,  I'l.- 

'Ii niliiisatiiiii   iiia\    Irail   In  I'liairiiialimi,   uliicli   explains  why.   if  tin 

I'liilli  piiiilin-iil  li.\  liiatini;  tin'  uliitr  nl'  an  i-u'ir  is  allnwcil  to  staiiil,  it  r:\i 
lint  ill'  auaiii  lifalrii  intn  a  I'lnlli.  tlir  allnimln  lia\  iiiir  urniif  nut  iif  snlulim 
liy  sill  I'ai'i'  I'liaiiulal  inn. 

An   inliTi'stiii'.'  plii'iinnii'iimi  i|i  iii'inlinu:  mi   tlw  .siirrai-i'  ti'iisim !it 

wki'ii  till'  piiilnplasmii-  cmitints  nl' a  I'iliafi-ii  int'iisiiriaii  is  pri'sscil  nut  i 
walii-  A  111'"  iin-iiiliiani'  I'mMis  mi  tin'  prntnplasiii  liccaiisi'  nt'  siirl'ai'i'  i-m 
ri-ntralimi  nf  all  mnsl  itumils  «  liii-li  Inwri-  siii-fai-c  I'lii'i'cy.     I!v  appiicali- 

"I'  'III'  piiiM-ipli'  nf  W'illaiil  (lil.l.s.  A.  I!.  .Ma-allliiii'-  i-iiii.-lui|.-s"tl:at  imt  

aiUni-ptinii.  as  lAJiiliiti'il  ill  a  rnllniilal  snliitimi.  liiit  also  the  local  ari-iiliinl 
tiniis  nf  niali'iial  nfti'ii  siTii  in  i-i'lU,  arc  assnciatcil  with  chaiiucs  in  mi 
iacc  fiici-n-y.  His  cniiclusimis  arc  liascil  lar<;cly  mi  microscopic  slinli 
of  various  forms  of  cell  cxhiliitiiiy:  ilitTcrcnt  i|c'_m-ccs  and  types  of  acti\  it 


I  III, I  III  I  w 


Ii7 


.ihil    illi:c'irnil|->l>     slllilic'il    liif    |Mi1ii>^illlM    li.V    riili.ill     III  \.lllilritiv       I'.y    Mirll 

,1    liiraiis    tlir    |iiil;i>»ilUII    stiiills    ihtcllM'    lihirU.       Ill     M'u:ftiilil lis.    Infill 

MflllMllllllillMS    'it'    |intil>»illlll    liriMir    <illl<r    IUMI     tllr    illllTliM't'    IlllM'.ll    till' 

.  Ii;if   mill    lilt'   rli!(iiii|'li>  I  riiuliiiiiiin:    |iiiil»   "T    till'    I'l  II    I  siiiriiiryiii  1    m- 

nil, I,. I'  ,1  pciili r  111 II  Miill   I'loiii  wliirli  liitrf  II  |inilnisM>n  i:i"\*>  mil 

I.I   liiriii   llii-   lii>t   still."'   ill   i-iiii.|ii!iiitiiiii.     '11 tviiiHlli    fnihi    tl 11. 

:i>  will  UN  till'  iii'riiiHiihiliiiii  lit'  |iiitiissiiiiii,  iii.'iv  111'  till'  ii'siilt  111'  II  liivv 
siiiriii'i-   ti'iisiiiii.      Ill    iiiiiri'lliiliir   Miiiiiiiii   iiiu;iiii>iiis.   siii'li    iiH    N'mtii'i'llii, 

liilli-ll  li'ss  piitiissillMI  l'>  IHi'MllI,  111  ill'_'  I'lillliliril  111  till'  liiiM'  iif  till'  rili;i. 
wliii'li  Miii'.'illiilii  liilii'Vis  iiiilii'iili's  tlnit  till'  sti  iH'tiin's  :.ii'  |iiiii|iii'iii  ms 
nil    iilltciiliii'   111'   liA\    sill  Tili'i'   trllsiiill. 

Ill  till'  I'i'lls  111'  hiiilicl-  iiniliiiils.  i|i|iiisits  111'  |iiit,is>,liiiii  nil'  jilsii  liiivili/i'il  ; 
ill   stiillti'il   llll|si-li',    fur  i'\lllii|i|i',   tlii'>    I'l'i-llf   111    11    /mil'  Ml    r.H-ll   rllil    lit'  till' 
.Imilily   I'l'I'l'Mi-lixr   luiinj  iiinl    iiiiliirili;it.'l>    iiiljiiri'iit    I"   tlii'   siirjly    ii'l'l-;ir 
tiM'  lijiml      Cliiiiiui's   ill   siiil,-iiM'  tiii>iiiii.  iissiii'jjitrii   witli   i'Iliii'.'i's   in    tlir 
iIinI  liliiitimi   III'  |iiil;issiiiiii.   iiii'   lii'lii-\  I'll    li,\    iiiiiiiv    In   lir    n's|iuMsilili'    I'nr 

lii'iM-lll.il litl'Mi-tiiill.       Ill    IMlM's   mill    liri'M'    I'llls,    |iiil;issilllii    is   nilHTIl 

li.'iti'ij  ilt  till'  lIXiill  illlii  ill  till'  sllll'ili'i's  lit'  '111'  I'i'lls  lliti'li'stim:  sllUlti's 
tiiiiis  iii'i'  iitTi'ii'il  til  I'Spliiiii  till'  ii'l;itiiiiislii|i  miiiiii','  i-Imii'.'is  in  siii  t'iicr 
tiiisiiiii  ,'it  till'  triliiiiiJltiiills  III'  iiMilis  I  syiiiipsi's,  ti'iliiiliiitiiilis  in  jjlalnl  illiil 
iiiiisi'li'  I'i'lls)  liiiiiiirlit  iiliiiiil  I'.v  till'  iiirvi'  iiii|iiilsi'  Mi'tiiii:  jis  ji  i-luiii'_'i'  in 

ili'i-tlii-    piilrlltiill,       Siul'ii •iiliilrlismiiiii    111'    piit.'lssiiiiii     liiis    !l|si>    ln'i'ii 

i.lisl'fMll    ilt    till'    lllllH'll    IlilllIlT    111'    '_'lilllil    fills    I  pmii'I'i'ilsl.    illlli    nil    lllC    111 

iiii'ii  suit'; Ill'  till'  I'i'lls  if  till'  li'iiiil  ti'liilli's.     Siii'li  iilisi'l'Viitiiiiis  iiiiiii'iiti' 

111  wliiil   wiiy  siirfiii'i'  ti'iisiiiii   iii:i.\    lir  ■■.■illiij  iiitn  pl:iy  t iilml  i-rlliiliir 

;Hti\  itii's.  Till'  (it'll!  is  iii'w  mill  nliuiist  iiiii'xplnii'il,  Init  tln-ii'  is  iilri'iiilx 
iiiiii'li  III  iiiilii'iiti'  lliiit  siiifiii'i'  riii'i'L'x  |ilii>s  ii  must  iiiipiirt..iit  inli'  ill  tin' 
iMi-rmiiiiii of  iiiiiin    I'l'lluliir  !ii-ti\  iliis. 


Conditions  That  Influence  or  Are  Influenced  by  Adsorption 

Electric  Changes,  I'n'siili's  nn'ii'  I'liiii-i'iiinitiuii,  nthir  fun-i's  ri'iin' 
i.lii  p!;iy  til  iissist  nr  ri'tmil  iulsiiiptiiiii.  Om'  i,f  tin'  nmst  iiiipiirtmit  "f 
iiisi'  is  I'lcftl'ii-iil.  .Must  snliils  ulii'li  piisiiit  lis  piiltii'li's  ill  ii  luiil  cm'i'.v 
.1  iii'siiitivc  cliiirifi'  iif  cli'i-tiii'ity.  snini'  :i  piisiti\i'  iiiic.  In  i'iiiifiirinit\  ,\  itii 
'111'  Williinl  (iililis  liiw,  ii  I'lnisliint  Ii'inli'iii'v  «  ill  I'sist  fur  tliis  frn  i-in'iy;- 
'i'  111'  ilMiiiiiislii'ii  liy  till'  iii'iitiiili/iitiim  nf  tlir  I'lri'lrif  i-limi-'i'.     Tliis  i-iiii 

Ill-   l.y   ili'positiiiii    1111    til''    iiili'if; <{   utinr    piirtii-li's   i-iirryinir   ;m 

•  iri'lric  i'li;ii'i:i'  "f  nppiisiti"  siiiii  nr  liy  till'  iii-ti'Ui  "f  tliiil  prcsi'iit  mi  imis. 
'  li.'iri'uiil  ill  siispi'iisimi  ill  Wiiti'i',  fur  iiistmn-i',  liiis  ii  iifjiitiM'  cliariri' 
li'  I'liljiiiilnl  ii'i'ii.  w  lii''!!  llils  i!  jiiisitivi'  i'li:iryi',  is  mlili'il  to  tlii'  snllltimi.  it 
\m11  lii'i'iiliif  iji'piisiti'il  nil  till'  I'lliiri-iiill.  lis  will  iilsn  till'  I'lltimis  nf  illl 
iliiil'/inili'    silll.       On    ill'i-niiilt     nf    I'li'rtrii'    ililsnl'ptinli,    liyi'stllfl's    illlli     liili' 


RA 


I'liVMriiniiMicM.  iivsis  (IK  I'livsioi.iMircAr.  I'noci'.ssi-s 


Milts  iii<'  iiilsiii  lii'il  iiiiii'li  rrioi'i'  fii'cly  tli.'iri  tlicy  woiilil  In-  if  the  prucfss 
ilf|irinl.i|  sdlily  oil  Mirl;i<-i'  i'iiikIciishI  iciii :  tliiit  is,  il'  ilic  (iililis  rorimila  is 
iisi'ij  to  (•;ilciil,it.'  \hf  ;i(lsi.i|)ii.ni,  it  will  uivc  values  that  arc  iiiiicji  licinw 
tllosc   ai-'-.i.illy    fiPlllKJ. 

It"  tlic  liissolvcl  siihstaiici'  aiiii  tlic  |iaitiflfs  lintli  liavc  tlif  same  clcctiic 
siu'ii.  ndsdiptioii  will  not  occur.  Filter  jiapcr,  for  example,  lias  a  iicffa 
tive  cliiirp-  and  can  not  tlieret'orc  adsorb  a  nc-rative  dvc  such  as  coiifro 
red  fas  shown  Iiy  tlie  dejitli  to  which  it  lieconics  stained);  whereas  it 
rcaijily  adsorlis  niu'Iit  hhic,  wiiicli  is  positively  charijvd.  If  tiie  neu'ativc 
eharirc  of  the  jiaiier  is  lowered,  it  lieeoiiies  capalile  of  .-idsorliinir  some  of 

the  ncf^ative  consro  red.    This  can  1 iTeeted  cither  hy  jilacin^'  the  jiapcr 

ill  alc(diol  or  liy  aiidiny;  iiior<_'anic  salts  iNaCM  to  the  water  witli  which 
it  is  ill  contact.  The  positive  (•liarycd  ions  of  .\a,  produced  liy  dissocia 
tioii,  neiilrali/.e  some  of  the  nc^jiativc  cliar-ic  on  the  paper,  and  allow  a 
cerlaiii  anioiiiil  of  ailsorptioii  of  the  neirative  (diarycd  eorijjo  red  to  oc 
eiir.      As   wo, ill!   lie   expected,   ;icids  ami   alkalies  are  capa'  le   of   ffreatly 

altering  the  electric  eharues  liy  the  11  and  (»ll  ions  whidi  they  trilmte 

Chemical  Forces.     If  the  nature  of  the  piiase  at  the  surface  of  whicli 

adsorption   oc-ciiis   is   such   that    ',t    can   enter   into   ehetiiieal    ( liiiiation 

with  the  siilistaiice  adsorlicd.  reactions  will  oc-iir  that  d.  not  oliey  the 
laws  of  iiijiss  a<'tion,  I'.y  adsorplion.  reactions  of  ;i  certain  type  iiia\  In' 
eiienuraired  over  other  reactions,  even  althoujih  the  necessary  reactiii'-' 
Slilist.-inecs   may    lie    present    in    the   solution    (specilie   adsorption^      The 

adsorliinir  siihstai itself  is  not,  however,  usii;ill_..    suseeplilile  of  chciii 

leal  c  haiiL'e  even  when  it  exists  as  very  minute  particles,  as  in  the  ea.se  ol 
colloi<lal  solutions.  \(  veitheli'.ss,  adsorjitioii  may  accelerate  ehemicil 
react  i(Jiis  liy  liriii-,:! iiu  tni;rilier  in  concentrated  form  sulistanccs  of  liiyli 
ch.iiiic-;il    reactivity.      In   siicli   cases  the  adsorlpliiu:  siilistance   itself  does 

not   enter  into  the  idii'iiiieal   reaction,  jiiid   can    lie   n veicd  at    the  eml 

ill  .111  iinchaii'^i  ,1  .•onditioii.  ||  ,i,-i.s  ;i..;  ;,  <.,ital\st  I  pa;re  7"_M.  .\s  w. 
shall  see  Ijiter.  en/ymes  ,iet  in  this  way  i.e.,  their  rate  of  reaction  i- 
coMlrollcd   liy  adsorption,* 

The   distinmiishinir    feature   of  such   a<!sorption    phenomena    is   that    .i 
eiir\i'  of  the   reaction    i  drawn    liy    plolliuM;  ninoiint    of  (diemical   cliani." 

•AT;,.lh,r   Ti,.|.„„,.  .,(   ,1„-  ,„|Im,,„,.  .1   M,ru..'  ,  .„  r«i    ,„,   tli,-  ,„„rs.-   .,f  .l„,i,„al   t..i,ti..ns   i^.  ... 
I-.  111.-  .1,  .,l.i,,i,v.-   iiiH.Hij.,-   .if  ,li.,r,..,il  .,M    SIM I,   i,.i,liMM.   .,.   Ihr   ,.v,.liiti,„,   Ml    l,,ri,n.    ,i,i,|    kK,,, 
tu.      SurLiir   I. lis,,. II    „MV    ;i!».,   ,.,ii.,     i,  i.ii.I.iIi.mi   ,,f   ,  li.  iiii,  .,1    ri  ;n  II..1IS.   a.-,    i~    ~<<-ii    in    lli,    Hirl.plr 
I. III.-     1..     III.-     s,^;,,,r,;,„,i,     ,,t     ,lil,„,,|.„,„,     „lii.l,     i:l.„l,i:il!y     .l.v.|..|.s     « li,„    .,     ^^      N.HII.,     s„l,il„,n 
""-"'    «'"'    •■'    ..     'I'l"'-''    li>'!>.,i,    -..Inlio,      Th.    .„.t.-„,.   ,,,„.„i,s   ,1,-ar.    l„-,.,iis>-    surf;.,,-   ,-„.-rKv    I 
pr.  v.-tit.  i|    til.     n  ,1.  In. II, 

.\ii    itn|...il.iiil    ,11,, I    ..I    s,i!!.i,,     i,,i.i,.,i    ,.„    ,|,.„ii.,,l    r,.i,li,.i,s    i,    .,N,i    M.fi    III    tin-    r.-I.Ui,in-l 
'"'"',•■"    .'.'.,"",■'    ""     ■■'■^"'l'"""    ....llr.i.m    ..I     L.M-.S    M„li„ii,     .,1    I...,,    .l,„„h,,,|    |,v    „|,„    V. .1,1,1,, 

li,|iii,ll.      Ilii-   lnwr   II,.'   siirl;,,-.-   t.-iis Ill,-   ur,.il,r  Ih,    s..l,il.ililv  ..l    th,    k^.      (l-iv>!,-ii  a,,,!   iiili..t 

r,r,-.    I, .r    ,-,;,•:■:!,.    :::■;;:    v:,,:,     -    i,i;,i,     ,„    .,l,,,t.,:.    I.,,| ,,i ,    ,„     ,,,i    ,l,.,i,    i,i    „:,„.,,        n„.    sl„ 

III.-    fllllllty    ,,f   att,in|.|,n,;    I,,    ,,,,A,i,t    ll„     I,,.,    ..(    j,,,,,  ^    |,,„„    n„„|,    „„  |,    .,.,    ,,, |    ,    .    i.,,^,,,,„„    ,|,, 

mill  ,,ils  ,,r  li),lr,„ail.,,iis 


(■<I|,I.(III>S 


69 


iij,';rmst  coricciitraliiiii  of  iciicliiiji  Milisliincfs  i  is  a  imiJilinla,  imliuutiiiK 
ili.il  till-  laws  of  mass  artiim  (pane  "J:!)  ai'c  iii>  l(iii(ri'i'  r<>lli>\vcil.  In 
(Milcr  ti)  lit'  alilf  t(i  (IctcnriiiK'  w  lictlicr  sonic  particular  pniccss  as,  for 
ixainplc.  a  fcriiicnlatioii  i)roccss,  or  tlic  alisorptioii  of  oxyjrcn  liy  Mood — 
is  caused  liy  adsoiptioii,  \vc  iiiusf  compare  its  ciiivcs,  const iiictcd  ac- 
cordiiii:  to  tlic  same  in-inciplcs.  with  tlic  typical  adsorption  curve.  A 
formula  may  lie  used  in  const ructiiif;  the  curves.  In  arrivinj;  at  this 
formula,  two  fai'ts  have  to  lie  renicmlicred:  (1)  As  adsorption  procei'ds 
and  less  anil  less  of  the  free  enerjry  on  the  adsorliiii";  surface  reniaiiis 
111  he  ncutrali/ed,  the  reaction  slows  olT,  until  equililiriuni  is  reached. 
The  more  dilute  the  solution,  the  i:reater  is  the  iiroiiortioii  of  its  con- 
Iciits  to  he  adsorhcd,  which  means  that  :  '  u  is  the  amount  of  sulistaiice 
adsorlied  from  a  certain  solution,  then,  from  a  solution  of  twice  that 
strenjtth.  somen  hat  less  than  "2  a  will  he  adsorhcd  i.e.,  a  multiplied 
hy  some  root  of  li.  Altlioii;.di  the  foriinda  is  one  lie|oiiy;iii^  to  the  class 
Uiiowii  as  paraliolic,  it  must  not  lie  a^sumcd  thai  every  reaction  wliidi 
liappens  to  fi've  such  a  ]iaraliolie  curve  ^sucli  as  the  comliinatioii  of  t  »j 
with  lieMioi;loliin  under  e.  rtain  eondilioiisl  (see  pa^e  :'.s:', )  nnist  he  one 
di  pendent  on  adsorption. 

II  must  he  understood  that  although  the  sulistaiice  that  is  removed 
froiii  a  solution  h\  adsorjitioii  is  no  Imiin'r  i-apalile  of  eont  riliutinu:  to  the 
conductivity  or  the  osmotic  pressure  of  the  solution,  it  is  nevertheless 
lint  so  (irmly  lixcil  that  it  can  not  lie  sci  free  afraiii  hy  pil.ely  mechanical 
1111  ans,  as  hy  constant  dilution  of  the  tliiid.  If  charcoal  which  has  ad- 
>iiilicd  su^ar  is  placed  in  a  dialy/er  made  of  memtirane  the  pores  of 
\\liich  allow  suu'ar  hut  not  charcoal  lo  pass  tluouudi.  the  siiirar  will 
■_riailuall\  he  removed  if  the  dial.\/ir  is  ininierscd  in  riinnini;  water.  A 
'irlain  eipiilil-iuni  exists  hetwccn  the  sulistance  adsorlied  and  the  same 
^ull^tan(•^■  still  remainih-i  in  solution.  If  the  latter  is  constantly  dimin- 
i^hiiiy  hy  ilial.\sis,  the  adsorption  eoinpouiid  must  lireak  down  to  main 
iMJii  the  cipiililirinm.  It  is  clear,  however,  that  ilie  process  of  icinoval 
\ull  he  extremely  slow.  The  aliilitv  of  adsorlied  suhstances  to  withstand 
iciiioval  hy  washiii!:  is  taken  advaiita<;c  of  li.\  nature  in  holdiin:  hack 
I'liiiilstulTs  in  the  soil. 


i 


Physiologic  Processes  Depending  on  Adsorption 

lii>tanccs  in  which  adsorption  inidouhtedly  pla\s  a  most  imimrtanl 
111  in  pliysioloiric  |)rocesses  are  as  follows: 

I.  The  action  of  en/.ymes  (see  paye  71  ). 

'-'.  The  ciiinhinatiiin  of  toxin  with  antitoxin  oi-cuin  accoidiii!^-  |o  the  laws 
'  iiiNoiplion  rather  than  tiiose  of  mass  action.  In  this  case  it  is  im- 
•Mtaiil  to  note  that  when  the  toxin  of  diphtheria  is  added  in  small  sue- 


7() 


I'llVSh  IM  lliMlcM.    liAsls   (IK    I'llYsiol.ddrrAf.    l'U(i(i;ssi;s 


(•(•sNi\f  <|ii.ifililii's  til  (li|ilit|i(ii;i  ;iiiti1ci.\iii,  iiioii'  loxiii  is  luiil  lalizcd  tliaii 
wlii'ii  till'  tiixiii  is  all  aijilcil  at  (nicc.  A  siniilai-  |iln'ii(inu'iiiiii  can  also  lir 
iilisiTM'il  liy  ailijiiii.'  lillci-  itapcr  tn  imuij:"  ftil,  limii'  iif  tile  iiiiriiu'lil  lioiiii; 
ailsnilMil  ulii'ii  till'  |ia|M  r  is  ailili'ij  in  sinall  iiiiantitics  tlian  wlicn  atlili'il 
iill  ill  "ii<'«'-  Till'  cNiilaiiatiuii  is  that  I'clalivt'ly  nimv  adsinption  dt'  a 
■.'ivin   siilistaiii'i i-uis    I'iium   a   ililiili'   tlian    t'l'mii   a   sttimi;  soliitiun    (el'. 

|ML!V    (lit  I. 

;>.  Tlif  si'iisiii/iim  nf  Icin-ocv  tcs  liv  upsnnins,  as  well  as  flic  siilisciiiicnt 
iii'_'csti(iii  lit'  l.ai-illi  l.y  the  scnsiti/cil  Icnciicytcs.  Imtli  of  wliidi  I'lillnw  tlic 
cniiisi'  III'  an  aiisi)ii)tiiin  fcactiuii. 

4.  Till'  ti'iiiiatiim  nf  ailsniptiuii  (•oiiipouinjs.  sm-li  as  flic  innrjranic  salts 
ami  pmlciiis  anil  tlic  cuiiiplcx  Ici'itliin  cninpiiniKls  tliat  can  lie  cxtfacfcil 
fi'iiiii  rn'j  \iilk  uf  liiaiii  tissue.  In  such  conipiiiiliils  the  laws  tij'  clicniical 
piupurliiin  nil  liin'j-ci'  hi'M.  ami  prupcrtics  may  lie  exhiliited  that  are  (|nitc 

ilifl'creiit  Ciiiii  those  (if  eiihcr  iii f  its  ciimpiincnfs.     When  yulU  of  cirir 

is  extfacleil  with  ether,  I'm-  example,  a  cnmpuniKl  dl'  lecithin  with  \ifelliii 
j:iies  info  siilufiiin,  altlmimh  vitellin  itself  is  quite  iiisnlnhle  in  efliei-.* 
Tliei-e  cm  lie  nu  duiilit  that  adsnipfiun  eiinipuuiids  uf  this  cliaiactcr  an 
vciy  .iliuiiil.iiit  in  liviipj  cells,  and  that  they  ,ire  e. instantly  lieinu'  fiiriiiei! 
and  linikeii  ddw  n. 


|.>     liiMii,;    -..1,11,,,,;-    ,,1    .ri:    ..n„,„im.    o.ii,:..    r..l    a,i,|    i,    .Iv,-    ,.,H,,1    f„.ti.-    in    tin-    |,r,-«,,nn-    .,f 

-",'^-"'-'     "'"'I'     '■>      iiI'i.i:m. ',-.,,■, I.      ,  .  ,ITIi,  .,11.,.,    ,.„,     I ,,     I.,     1„      ,,.:,,, |     ,,,     ,,     ^,,1      -,     »,||',,, 

■■"■I    :.    i.T.,.1    i,,l|,,„h,l    |.,rlj.l,.       l-l,,     :itl,;.,iu>     I,..,-.,    l„.l,|i,,K    til,     ,,:„iul,-    l,,k',tli,r    iv    il.oiri.-    ,■ 


CIIAITKR  IX 
KHK.MKNTS,  OK   lAZYMIlS 

( ii t'  Itii'  niiist  strikiiiH:  (lfvclii|)iiit'iits  i>\'  iiiodcrii  rcsc:ii-"li  in  liinclii'iii 

isirx  (MiiH'ci-iis  llic  iijiliirc  i>(  ciizyiiii'  ac-tiun.  So  icinarkalilc  arr  iiiaii.v  nt 
ilii'  I'acts  that  liavc  Ipccii  l)rciiij;lit  to  li«:lit  lliat  it  <-aii  not  tail  to  interest 
(•\eiy  one  en^Mi:e<l  in  the  stinly  of  life  jilienoniena  whatever  the  natllie 
of  that  stnily  iriay  he  to  know  sometliiii','  of  the  main  (iiiestions  at 
present  oeenjiy iiiir  the  attention  of  investi'.'ators  in  this  liehi.  In  this 
eiiaiiter  a  lirief  stirve,\  will  he  iiiven  of  some  of  the>i'  i|Ueslions;  no  at- 
leiiipt  will  lie  iriade  at  eoni|ileteness,  ami  only  whire  neeessary  for  the 
sake  of  example  will  reference  lie  made  to  indixidual  types  of  en/.\  me 
ai-tion. 

The  diseoveiy  liy  Huehner  that  an  enzyme  can  lie  expressed  from  \ cast 
ei'lls  which  is  eapahle  of  instantly  lirintrin-j;  ali<i\it  the  alcoholic  fernien- 
lation  of  dextrose  solutions  lias  lieeii  res|ionsil.le  for  a  L'leal  deal  of  the 
imideiii  advance.  Formerly,  yeast  cells  were  lielieved  to  lirinu'  almnt 
.ilcoholic  fermentation  as  a  resnlt  of  their  p:rowth:  it  was  lielieved  to  lie 
a  life  iihenomenon,  or  "vital  process."  Now  we  know  that  yeast  cells 
pnidiice  an  intracellular  feriin'nt  or  endo-eiizyme*  to  which  its  sncroelastic 
piiiperties  are  due  and  wlii(di  can  act  apart  from  the  cells  that  produi'c  it. 
Il  is  no  ^reat  stretch  of  imagination  to  think  of  all  chemical  reactions 
ni- dialed  liy  cellular  ai'tivity  as  due  to  a  similar  mechanism,  and  this  thoULdit 
has  h'd  to  the  hypothesis  that  all  processes  of  inti'rmcdiary  metaholism  in 
the  animal  ami  plant  are  cailseil  hy  en/.ynu'  action.  Hefore  I'.uchner's 
.lay  w<'  knew  only  of  the  e\l  racellillar  en/\  nies  ismdi,  for  example,  as 
the  di<.'estive  ferments),  that  is  to  say,  of  en/ymes,  proijncid  indce<l  liy 
.ells,  hut  sei-reted   from  them  and  actinii  outside  thi'ir  protoplasm;  now 

u.'  must   I yrnize  intracellular  enzymes  acting  where  they  are  produced. 

"  ihe  protoplasm  of  the  cell.  I'.ut  we  must  not  permit  this  conception  to 
carry  us  too  far.  Without  furtlu'r  invotiiratiou  we  must  not  imairine 
ill'. I  Ihe  riddle  of  life  is  thus  solved. 

As  an  example  of  tlu>  role  which  extra-  and  intracellular  enzymes  are 

-iipposed  to  play  in  the  animal  momy  max   lie  cited  the  metaliolism  of 

:'!'>icin.  Proteolytic  enzyiiu's  are  very  widely  dist  rilmted  in  the  acti\e 
'.^Niirs  of  the  animal  and  i)lant.     H.\   their  aucncy  in  animal  life,  the  < 

I  il.    i.nn-   "1".  rim  lit"   ;ni'i   'Vn/vnu"  .tri    w  iimiu  m.-u-.    I';:!    III.    I.;n<  i    t>   |i.i.i.ii.l.     .i-   flu     i 

•■  111.    I-Tiii.  r  t.,  I,,'  n~.-.|  .,-  ilii"  \.  il.. 


<-  I'ln  SM  HI  iiiMli  \i.   iiAsis  (PK   i'ii\  s'(U.(M;ir\r,   I'I|(H'|;ssi;s 

l>lrx  |)i-i)t('iii   iiKilci'iilc  is  split   ii|i  111  ii-nilrr  it  alisuiliiililc  ri'iiiii  tlir  ililcs 
tirii',  iilld  till'  tissues  .llililiil'l-iiiti'   lldlil   the  lilimil  tllosc  iif  tlic  (Iciri-iuliltinil 

lU'iMJiii'ts  tli.it   tlii'v   rii|iiii('   fur  III iistnictiiiii   (if  protiipliisiii,   uliicli. 

liiti'i-,   tlii-y    iliiMiiii|iosc   sii   iis   to    iilili/c   tlic   i'iiiT<iy    «]iicli    tllc   oi'Killlislii 
(ii'iiiiiiiijs.     All  tlicsc  piiiiM'sscs  iiic  li('lic\<'i|  til  lie  tlic  work  of  I'li/yiiics 

The  Nature  of  Enzyme  Action 

'riic  fli.iiiiri's  liinimlit  Illiniil  liy  cli/yiiii's  i-;iil  also  ln'  aci-omiilisllcd  li.\ 
onliiiaiy  rlii'iiiii-al  iihmiis,  Imt  tlirsc  liavc  ot'tfu  to  l)c  of  a  very  ciici-fjctic 
iiatuii'  to  acroiiiplisli  what  tlic  cii/.yiiif  can  so  (piickly  and  (piictly 
|)cr  I'orni. 

It  is  till'  lUstoMi  to  iciraiil  cii/yiiii's  as  rnldhisls.  A  catalvst  is  a  suli 
stance  uliieli  ai'celcrates  lor  letanlsi  a  eliciiiieal  reaction  uliich  in  its 
aliseiice  could  proi-i'id  at  a  ditVeii'iit  i  usually  slower)  |)ace.  The  action 
of  catalvsts  lias  liecii  aptly  likened  to  tliat  of  a  Inlii'icant.  A  weijilit 
lilaccd  at  till'  top  of  an  iiicliiied  plane,  so  lield  that  the  wciirlit  only  sloulv 
slips  doxMi.  lias  its  velocity  yicaliv  increased  if  its  uiulei'  surface  he 
oiled.  The  oil  accelerates  the  action  hut  does  not  alTect  the  ultimate 
result.  Catalysts  do  not  condiiiie  with  'he  final  products  of  the  reaction, 
these  lieiiii_>-,  as  a  rule,  the  same  as  they  would  ha\c  licen  had  no  eatalvst 
licen  added.  Another  characteristic  is  the  ticniendous  aunnuit  of  eheiii 
ical  chanLie  which  even  a  trace  of  catalyst  can  induce.  Tlu't'c  are  iiiain 
examples  of  catalvsts  in  the  inoriranie  world,  anione-  which  may  he  cited 
the  aelion  of  spouirv  plalinum  oii  liydroL;cn  pcioxide.  This  sulistaui-e 
normally  tends  to  decouiiiose  into  watei'  and  o\vi,'en,  hut  if  a  siiiali 
amount  of  spoir.;v    platinum   is  added   to  it.  the  decoin|)osition   is  '_M-eallv 

i eleiated:    II   (I  1 1  .<)      (» 

A  very  iriioij  cNample  of  the  action  of  an  inoi'iranic  catalyst  is  that  o' 
the  hvdriejeii  iiiii  oil  caiie  siiu'ar.  or  other  disaecliarides,  in  the  prcseuei 
"'  "■•I'ei-.  It  aeci'lirates  the  hydi'olysis.  Cane  suirar  solution  at  rooii; 
temperalure  docs  iiot  indeed,  in  sterile  solution,  undeiLro  jiiiy  .-ippi-ccialil' 
hydrolysis,  hut  at    luii     ( '.  ii  .hies,  which  lea.ls  us  to  helieve  that,  thoUi;! 

ii'-'l iaiile.  the  adion   .ilso  oeeiirs  ;it    i-ooiii   tciiipcratur-c.     I'.v   addiu'j 

•■I  vdroehloiic  acid,  or  other  acid   not    liav  inj.'  an   oxidizinir  elTe.' 

on  su^ar.  we  !,;ri'all>    accelerate  liie  hvdrolysis  liecanse  of  the  hydroirr' 

ions  ines.nt    in   the  ;ii-id  solution.     Withii rtain   limits  the  rate  of  li\ 

diolvsis  is  propoi  iion;il  to  the  amount  of  catalyst  prrspiit. 

l-ji/yiius.    likr   otiiir  catalvsts,    pioduce   their   ai'tioil    when    present    i 

\''l'>   small  i lints  i  e.  ._r  ,  siicrase  i-an  hydroly/c  2()(»,()(l()  times  its  weii;! 

of  cane  siiiiar;  diastase  can  convert  starch  to  snv'iir  in  a  dilution  . 
1  l,ililil.(lliii  ,'111, 1  till. I,,  is  a  distinct  rci;ilions|ii|,  lictvveeu  the  amount  " 
.■n/vhic  iireseiil   and  the   rate  of  th,.   retictioii.     The  finid   product   of  tl. 


KKRMKNTS,   OK    I'.NZVMKS 


7:j 


icjictiim  is,  liowcvcr.  (lie  siiiiic  at  wliatcvt-r  into  it  pincccils,  iuid  the 
riizvinc  (Iocs  not  iippt'iir  in  tlic  tiiial  imnlucls.  Ahmy  fii/.vnics  such  as 
iliiistasc  can  lie  Idiind  \iiialtcic(l  in  anmurit  after  tlicy  liavc  cmniilctcil 
their  action.  This  is  (leteniiincd  hv  addinu'  a  fresii  siijiply  ol'  siihstiate 
I  that  is,  of  material  to  lie  acted  oni.  when  the  enz\  luic  action  proceeds 
.I'.'ain  in  tlie  usual  \va\ .  The  sanu-  is  no  doiilit  true  for  all  cn/.\  nies, 
ihoujrh  as  yet  it  can  actually  he  pinvcd  for  only  a  feu  of  thcui.  Kii/>  iiies, 
therefore,  nniy  he  defined  as  catalysts  iiroduced  liy  liviii>;  or'.'anisnis. 

The  Properties  of  Enzymes 

Altliouuli  en/ynies  ale  examples  of  catalysts,  they  eshihit  many  jiropcr- 
tics  that  ai)i>ear  to  ditVer  from  those  of  inoruMnic  eatal>s1s.  It  will, 
llieret'ore,  l)e  advisalile  in  considerin'^  each  quality  t<i  compare  it  in 
catalysts  ami  en/ymes,  for  hy  this  method  a  much  clearer  conception  of 
the  nature  of  enz,\ me  action  can  he  >_'ained  (  I'.a.\  liss' '  i.  Those  ])ropertics 
that  are  strictly  peculiar  to  <'n/,\  lucs  we  shall  consider  later. 

I.  Most  tii:!iiiiii  iii'i  rniiiirl.iiJilii  spirifn'  in  lliilr  dfliau,  irliirnm  iiior- 
ijiniir  cdliiliislx  an  ririi  nnicli  hss  si>.  Thus,  in  the  case  of  the  enzymes 
«hieli  hrin<r  ahoii!  inversion  of  disaecharidi's.  this  specilicity  is  clearly 
shown.  There  is  a  spei-ial  enzyme  for  eai'h  of  the  three  ilisaechnrides 
maltose,  lactose  and  cane  suirar  and  one  of  these  can  not  i-eplaee 
another. 

Stii;  more  stril-ini;l.\    is  this  s| ilicit.x    of  cnz.x  nu>  ai-tion  demonstrated 

ill  the  fact  that  certain  enzyim's,  sui'h  as  z,\  niiise  (expressed  from  yejistK 
will  act  only  on  hodies  havini:  a  certain  conliixuration,  that  is.  havinsr 
their  side  (diaiiis  !irran<;iMl  in  a  certain  \va.\ .  Thus,  there  iire  two  vjirie- 
ties  of  dextrose  (u  and  fiK  which  differ  from  each  other  solely  in  the 
t'ai't  that  the  side  chains  ari-  tirraniT'd  in  ditTerent  positions  with  rela- 
tion to  the  central  (diain  of  carhon  iitmiis.  This  form  of  isomerism  is 
-ailed  stereoisomerism  lieeause  the  two  hodies  rotate  the  jilanc  of  polar 
ize.l  liirht  to  an  eipi.-d  dcirrei'  in  oi)posite  directions.  Zymase  iicts  on  one 
nf  these  hut  not  on  the  other,  and  there  are  iiinumerahle  cxami)les  of  the 
^iiiiic  kind.  Indeed,  of  all  hodies  that  exist  in  two  stereoisomers  only 
one  is  found  in  li\  int'  cells  and  it  is  on  this  variet.v  alone  that  the  eorv  mes 
ill  animals  can  act.  A  similar  specilicity  exists  hetween  certain  ^\■  and 
'hi'ir  pharmai'olofric  ;iction. 

Spicilicity  of  action  is  explained  hy  supposinsr  that  a  union  occurs 
i"l^^cen  the  sid)strate  and  the  enzyTue,  and  for  this  union  to  take 
ii'.iee  the  enzyme  must  jiosscss  a  eontiu'iiratioii  which  corresponds  accu- 
i.itrly  with  that  of  the  stilistrate.  The  process  has  heen  compared  to  a 
'•'"■k  and  key;  the  key  nitl.st  he  shaped  to  tit  the  lock,  or  it  can  not 
"i'lrate.     The  specilicity  <loes  not,   however,   in   itself  disprove  the  i-lose 


H 


74 


l'IIV>l(  (H  III  MUM.    l:\sl.s    OK    ni  VSIul.udir.M.    I'HIHI>SIS 


nliitidiisliip    lictwr.ii    cii/yiiu's   iiiiil    iiiiii'jiinic   cntiilN  sis.    (i<v  <iii    lln-   iiiir 
li.'iiicl  tIl.Tc  lire  M'Nciiil  ni/viiics  wliii-li  i|i>  imt  cNliiliil   tliis  pmiifrly,  :inc| 
nil    llic   iillicr.   llicTt'   ;irc'    iiinrLraiiic   c.-itiilN  >ls    whidi    ,1,,.      l'„v   (■xiiiii|>Ic, 
li|)iisc.  llic  t';il  splillilii:  (li/viiic  III'  |i:iii<M,';ilic  Juice,  (liciiiiipost's  imt   (Hil> 
IiiIn  Imt   til  ;i  yii'jiI.T  ur  less  dfoici'  ,i   iiiiiiilii'i-  iif  Imdii's  iil'  llic  s.iliic  i:cm 
•'I'll    liiiilil    (c^lc|■^!.    iiiiil    t\  r.iNiiijisc    ciui    ili'i'iiiiiposc,    iiof    tyiosiii    iilidic, 
Imt   ;ill   plii'iiiil   cciiiipdiiMijs.      r<p|i\ciscly,  the   li.\  iliuircii    ion      Id  llic   ))ic> 
clicc  of  wliicli  iiciils  iiwc  llicMi'  i-iit,-||yt  ic   pdWciN      ciiil  ilcciiillpcisc  llic  (iliji 
iijiry   caters  illijil    is.  ni'  jn-iik  ccmt.-rniiii','  tiic  ciiilMiNyj   ,,]■  COOlI   f,'rciiip 
Imt    it    h.lN  no  ••irljuii   nil    llic  Milplidliic  cslcls.      jidwcvcr,  c|i/,\  iiics  lire   n-l 
t:iiiily  ijiiicli  more  spceili.^  in  ilicir  iiclioii  lliiiii  iiioririiiiic  ciitalyNls. 

-.  'I  I  iii/ii  nil  iiri  iliiis  iifil  nilhdiiri  raliiliixis  mnl  iimiiiii  iicllnii  in  llu 
s'liiii  irmi.  As  ijie  tciiiperatiiic  is  rais.il  in  the  .■.isc  of  inor^'aiiic  catalysis, 
the  reaction  Im'coimcs  alioiit  doiilpjcd  in  rapiiiity  for  I'licli  risi.  of  10  ('.. 
wlicrcMs  in  the  easc>  of  eii/yincs  it  li.'coin.'s  increased  up  to  a  certain  iii>li- 
mil, II  I  <  mill  rut  in; ,  lie,\oiid  wliidi,  as  tlie  teiiipcrat  ure  rises,  tlic  reactjcui  i-- 
lirst   slow  I'll  ami   tlicii   disappears  aitoycllicr, 

Tliis  peculiarity  of  eii/yiiio  as  compared  «illi  iiiorL'.'iiiic  catalysis  need 
not  in  itself  disprove  ihe  aiialo.^y  lictueen  llic  two,  liecailsc  cii/yiiies  d" 
not  form  irue,  Imt  colloidal  solutions.  Colloidal  solutions,  as  we  Iia\i' 
seen,  arc  r.'allx  line  suspeiision>  of  nil  rainicroscopic  jiarticlcs ;  llicrc  is  \\« 
splittini;  into  ioii>  of  the  dissolved  Milistancc,  as  is  tlic  case  xvilli  tiMie 
iinolcciilari  solutions.  Imt  tlic  colloid  is  suspended  in  ||u-  water  nv  otlur 
~^''l\|'ii'  '"  l'"Mii  a  lieleroireiieoii.s  system  ipa^-C  •">  I  ) .  oil  wliicli  account 
the  siirfai-e  area  oi'  the  iiiciisl riiiiiii  is  ciiiu-iiioilsly  incrcjiscd.  Rise  ii 
tempiM-atiiie  alters  the  estciit  of  the  siiriacc  area,  and  therchy  intro 
duces  an   inllueiicc   which   proLrressi\  ely  opposes  calalvsis. 

.MthiMi'.di  iiioriranic  catalysts  in  molecular  solution  show  no  opliiiiun 
teniperatiirc  Imt  increase  in  acti\it\  in  proportion  as  the  teiiiperature  i 
raised,  iiiori^anic  colloidal  catal.\sls  iiia.\  show  an  optimum  feiiipcratui' 
Thus,  spoin.'y  platinum  shows  an  optimum  temperature  in  its  action  on 
mix'iiri'  of  h.\drnueii  aihl  ox\i;en.  |t  lias  therc't'orc  heen  suij^'cstcd  tlm 
It   IS  liccause  tlic\    are  colloiiK  that   cn/yines  cxhihit   this  pr(>|)crty. 

:!.  I  iiiininiii,  ,-ii!,ihists  In  i/iii  ii/h/  ,;n-ni  tin  rnulinn  I,,  ,i  fiirlhir  .s/.e 
llmii  Hull  all, mini  hii  llu  nrlluii  ,il  iir.miiis.  j-'or  cxaiiipli',  aciil  lircak 
ilow  II  the  protein  molecule  much  moll'  coiii|ilciely  than  del  the  iiroteolMl 
enzymes.  'I'his  diir.'rciicc  is  p.M'haps  .'Nplaiiied  hy  the  fact  that  eiizym 
are  retanlcd  in  their  acti\  itii's  w  hen  there  comes  to  lie  a  certain  ai'ciiiii' 
latioii  of  the  procliicts  of  the  reaction  iircsciit.  The  liiial  sfa<:cs  in  ti 
reaction  may  hecome  so  slow  as  to  lie  almost  inaiiprccialilc.  This  d- 
••rcasi'  in  acti\il,\  is  parllv  due  to  a  union  lictwccn  the  cnzyiiie  ;iiid  ti 
prndiicts  of  its  activity. 


I'l.liMIAIS.    III!    lA/.YMI.S 


75 


t.  '//((  '•iliicilij  ntuslititl  ill  till  iiisi  of  iiiiiriinii'ir  nilulitsls  n  niiiliis  nil- 
■  liiiiiiiiil  llinii'iildiiit  till  riiiiliiiii.  irhiri'ts  in  lln  nisi  nl  i  ii:iiiiii i^^t  hi  ■ 
"/(IIS-  lilhir  liss  iir  t/ridii  r  na  tin  iimn  ss  ininnih.  Wlicii  ii  siiliNtiiiicc  is 
.liiiMU'i'd  li>  (Mtiil.v  lif  jictiun.  It  is.  of  chuim',  cciiistanlly  liciiii;  diiiiiiiislicil 
III  iMPiii'ciilijii  lull  so  tli:it  Ifss  iind  li'ss  ol'  i1  rtiiiirnis  to  lie  jii-ti'd  on.  Tliis 
iiiililics  tliJit  tlicri'  lire  IVwcr  inoliciilcs  picsciit  t'or  tlu'  .siiiiii'  iiliiotiiit  ot" 
.iliilyst  to  net  on  iiml  consi'iiuciitlv  lli.-il  tlic  imionnt  clinnjii'd  in  ii  unit 
nl'  tiiiii'  is  ])roi:rcssi\cly  less.  At  ;in.\  iiionii'iit.  liicicfori-,  llii'  lute  <d" 
i;iliii\sis  will  lit'  |>r<>i>oition;d  to  tin'  iiiiioiint  of  Milistiiiii-c  ( sniistniti' ' 
I.  ft.  To  iiiidcrstiind  this  we  iiiiist  refer  liiiek  to  wliiit  we  liiive  leiirned  iilmnt 
iiiMss  aetioll.  If  we  supimse  tllilt  two  snlistilliees  A  iillcl  15  reilet  to  follil 
iwo  otiier  snlistilliees  ('  ;iiid  |),  then,  li.\  the  liiu  of  mass  nelion,  the  reai-- 
limi  vv  ill  not  ir<>  on  to  euiiipleiion  Imt  w  ill  stop  w  hen  :i  eertiiin  ei|nililiriiini 
is      levelled.        'I'lli'      re;n'tioii      ejill      lie      represented      li.\       the      ei|nation 

.\     I'.  F^C     I),   wliieh   means  that    it    pr Is  at   a    rale   propoitjonal    t" 

llie  reaetilijr  iiioleenles.  In  smile  cases  tliis  reai'tion  L'oes  <in  nnlil  eitlief 
A  or  11  has  jiraetieall.v  cjisappeaied  ithat  is,  the  ei{nililiiiniii  point  is  vei-.v 
mar  the  rii:ht  of  the  «'ipialioni.  as  is  tiie  ease  in  the  inversion  of  cane 
>iiL'ar: 

<',,H,, (),,    irj)     (",  ii,,(>,    < ,  ir„o, 

'i'akin!;  place  as  it  does  in  an  excels  of  water,  and  tlieie  lieiiiy;  \el'.\ 
liille  tendency  I'or  this  reaction  to  ;;ii  in  tlic  o|ipositc  dirccti'ni  (ef.  re- 
\ilsililr  actioiit  I  pa'_'e  'Jo  1 ,  the  olll.\  tliiiitr  which  will  inllllellce  its 
velocity  is  the  eonceiitration  of  cane  sii'-'ar:  in  other  words,  tlic  velocity 
I.''  the  reaction  at  any  nioinent  will  depend  solely  on  the  eoiicentratioii, 
• '.  of  the  material  still  h'ft  niitleeomposcd.  This  can  lie  detei milled  li.v 
means  of  an  ei(iiatioii.* 

The  value  of  such  an  i'i|uation  is  that  it  uives  ns  a  (iuilie  K.  icpieseiit- 
inu'  Ihe  amount  of  inversion  that  would  occur  in  each  unit  of  time  if  the 
iMiie  siiuar  were  Uept  in  constant  concentration.  Wlien,  for  exaiiiplt", 
I'  is  stateil  tiiat  K  for  a  particular  strenuth  of  acid  actiim  on  cane  sny:ar 
^^ll|lioll  is  O.Od'J,  this  means  that  when  \oluiiie,  conc<'iitratioii  of  ;icid  ami 

■h    ■.    I.,     til,     .111;. 1, ml    "f   -l-Mr    iiiv.ll..!    in    li    '.     I      i:u\    if    ^^  ,     M-i     ;i    "l:"'.     ..ill.,!    :i   .    ■r-'.int    ih'l    I" 
'.--    Ihi-    111  i.i;i:ii.nl,.l    lal.-    i.|'    llii-    1..1.I11II    Iwlii.h    will    llur.f..rc     1..     .lill.nnl    l..r    .lniii.iit    ri:n" 

-'.   ti..!i  Kv'        loit    i'    ..-111    'i..t    i.f  ill.-    '.I'lH    .il   .tny    X\\  >   ...n-..nliv.     [..ri...'^    ..1    liin.'.    t.if.r.i^L- 

I.  .  t    .1;  I-  i;..iin.  ..■:  .■.iiliiiii.iiil>.     Tlii-  11  11.1.  1-  il  ■ii.'.  --;in   I..  11-.    III.'  i...m1' I   !!■. T,  r.  iiti.il 


ls>'.        I'll.     -ii;;.    ,1    inl!.,i!,.    ill,.'     tlu-    r.':..li,.li    1-    .1    .■,.r-l;iiilK     i  li:.I!f:il 


'li;iT    o\    :in.!    „i    !.|,.,-.nl    -.i.li    inlinii.ly    >i;,,ill    r'lv,,  ,i!iit^    ih.il    ;!i.\'    tun    i',,l    !,.■    ni.M^iin 
i-   "1    in!vL:r,i:i..ii.    l,-,\\.Mr.    il    cm    I,;-    -l|,.wii    tli.it    ttl.     .tl-.vr    (.ji'ati.ii    ii.,iv    1,.-    uiill.ii: 


r.     1  I  .■ 

■ '■"iliiii,'    11-    1.1    lii'.l    ih,-    \.,l.,i,-   ,,|-    K    II',    I',    li.  iiii;    tl;f    i  ,.iu  1  iilr.iti,.ii>   •<    .■,iiii-    -ii,l...t;-    .it    11:.- 
■  ■    *i  ,    'i' ..  1 . 

\'  V    li\..  .|.l.riniii.iii..i  ^  .luiiin,'  till     ...'II-.-  ..f   ill.-    1,  .-iili,.ii    1.-11.    1..-   11-1.1    l.T   .   ,1.    il  iliiii.'    K.      'I'l 

.'I, ,11-    -iiii.lv    .,i,lv    I..   i.iM.-    II.    vviM.li    1,111    ..lu-    •.iil.-l.iii.  .     --    ■  li.iii!;ii.«    111 -i...!.-.  ul.ir    i..iili.,ni. 

.  .1    1»..   -.|l.  t.tiu,-    .-111.    in\..lv..l.    III.-    i.|.;ati..tl    '^    ti!,-u-    .-,,iiii,li.  ;iU  ,1. 


Pffn 


fi 


nivsii  (11  III  MUM.    liAsIS   (IK    IIIVslol.iMiKAI.    I'HOIISSIIH 


I 


ti'iiiiiiTiiliiri'  .III'  i-..ii>t.iiil    in   it    •_'i;iiii  liiiilfciiliir  siiliilimi  of  su«iir,   d.lMIL' 
Kiiiiii;^ii|iciilc   i,(  swiTtw    wi.iiM    lif    iiivcitcd    the    first    iiiiiiiitc   niKl   (».(MIJ 
U'liini  I'Jii'li  Mircc.'.liii'.'  liiiniitc,  |i|ci\  iilcil   uc  coiiltl   kerp  the  siiliitiiiii  ciiii 
Nl.-intiv  ii  m-.iiii  iihiIiciiImi-  i.iir.  tluit   is,  i)i(i\  idol  we  cDuld  iuM  sii(.'iir  Jiisl 
iis  (|iiicldv  MS  it  inciPiiics  invfiti'd. 

At  first  siiTJit  it  fii.-iy  ,i|)|ii;ir  of  littli'  |ir;ictii'ai  iiiiportancc  to  (li'tiTiniiif 

'^-      III   "III'  I'lisciit   disciissiiM iiciTiiiiijr  till-   iiatii?-!'  of  (•ii/yiiic  iii'tioii. 

it  is  lioui'MT  of  i.'i,';it  viiliic  for,  w  licicas  witli  irior'.'aiiic  catalysis  l\  i^ 
really  of  coiistaiil  \mIiic,  witli  cii/ynic  a.-tion  it  is  not  so.  Tims,  mImmi 
caric  simar  is  iiixi'itcd  liy  sm-iasc  an  cii/ynic  prcsi'iit  in  tlio  inti-stiirr 
an. I  in  yeast  tlie  roiislant  uradnally  rises;  for  most  otlxr  iiiiiinoleciilar 
reaetioiis  mediated  li\  eiizyiiies  it  u'radiially  fails;  for  example,  flio  aetion 
of  trypsin   on   jiroteins. 

Where  Ihei-e  is  a  ureat  excess  of  slilistance  to  lie  acted  on.  in  coinl>ari 
son  with  ttie  ainoiint  of  eii/.v  me  present,  it  will  lie  found  lliat  a  nunc 
constant  value  than  K  is  ohtaine.l  when  wc  compute  the  ahsojiite  anniuni 

of   sulistaiice   <|i niposed    in    a    jiixiMi    time.      In    such    a    c;ise,    too,   tin' 

amount  of  chanue  in  a  i^ivcn  time  will  he  proportional  to  the  aiiioiiiit  ot 
en/ynie  pi-csenl,  indii-al iin.'  that  some  soft  of  eomliination  lii'twoen  en 
/.\  me  and   siihstrale  must    Im>  the   first   step   in    the    fermentative   |irocess 

This   fac-t    has   heen    noiicc'd    hy    us   ii i tioii    with   the  hydrolysis  of 

yly<-o'_'en  in  the  jixfr.  When  then'  is  an  excess  of  i;l.\-eo<rcn  present,  fin 
amounts  uhich  disappear  in  e<pi;ii   intervals  of  time  after  death  are  the 

sano';   when,  on   the  contrary,  tli.'ie   is  not    much   trl.v  ••ou'en,   the  a um 

^^hie||  disappears  -iradualiy  d. •.•lines.  I. ill.  if  K  I >iii|nit.>.|  |,y  the  alio\c 

e.platioii.   it    is  .-oiistant. 

'P.i  niaUe  these    faints  .'li'ar  it    may   he   well    to   pause    foi'  ji    nioinent    t" 

coiisiiler  an  illiisl  rali.m.     Th nilithms  ..litainin<r  w  lii-ii  there  is  a  lari;- 

.x.-.'ss  .jf  siiiistrati'  oMM    .'ii/.v  111.'  iii;iy   III"  ipareil   to   those  '„'overniir 

the  removal  ..f  a  pile  of  l,ri,-Ks  from  one  pla.'c  t.i  another  hy  a  numlii'r  ..i 
'ii'''i      'I'll''  I'll''  "(  Ini.-ks  r.'presents  the  sulistrati';  the  men.  the  .•ii/,ymr 
If  ea.-h    man    w.irks   up  I.,   his  cap;i.-iiy,   it    is    plain    that    the    niiiidier  ..■ 
hri.-ks  traiisferr,.,!   in  a   yiven  time  will   not   .li-p.^nd  at  all  on   the  size  .  ■ 
the  ]iil.'  to  hi'  transi'errid.     When.  In.ueM-r,  the  pile  of  hricks  irets  sma!' 
thoiii.'h  the  sam.'  niimlH'r  of  men  cnntinue  to  work   tin-  niiiiilicr  of  hri.-k 
transt'erred  in  a  -.'iv.ii  tim.'  falls  .ifV.  liecause  the  men  interfere  witli  ut., 
iinother's  acti\  ities  in  seciirinir  their  loads  from  the  pile.     When  a  simi!,. 
sta<re  is  ariived  at   in  en/.\  m.'  processes,  we  have  to  use  the  velocity  cm 
staiit  to  shnu  iiow   niiich  wmk  could  lie  done  li.\   th.'  enzyme  if  the  jimoin 
of  sllhstrate  were  inaintaiiKMi  of  constant  amount. 

Jii   the   larir.'   volm f   recent    w.irk   whi.'li   has   lieen   done   with   th 

<ili,ii'ct  of  dis.-.iM'riiiir  the  .'ausc  of  these  variations   in   the  velocitv   en; 


KIKMINTS.   OK    l;\/.YMi;s 


77 


stiUil  ill  till'  ''asf  (if  rli/..Mlirs,  rolir  ilMpurtiilil  nilnlitiolis  liilNf  liccil  n-cd^- 
iii/i'd;  (1  1  I'cvci^iliility  :  I'i  i  uriiiliuil  <|rsliurti(ili  (if  tlic  cii/viiu';  C!)  coiii- 
liiiiiiliiiii  III"  tl ii/viiii'  uitli   |)ri)i|iifls  nl"  tlic  ii'iictinii;  (4)   iiutuciilulyNis. 

(•!'  flit'sf  I'diir  inlliH'iict's  tlic  mily  oiic  wliicli  cniilil  !»■  licld  m iiiitii1il(> 

tnr  Jill  iinTi'iisc  ill  tlir  ;iclivily  i<(  tin'  cli/yiiK-  is  iiiiliiiiildhisis:  in  this 
|p|(H'css  till'  rii/yiiic  liv  its  iirtioll  |i|<nllli'i'S  slllistillircs  wliicli  iiitcMsify 
iis  own  ii('ti\it>.  In  smiic  cases  at  Iciist  Cur  cMiinpli',  tlic  iicfion  of 
iiiM'itiisc  nil  cane  siiirar  tlicsc  arc  aciil  Imilics.  a  iiiodcralc  increase  in 
iiciility  t'avcii'ini,'  the  action  of  this  enzymo. 

'riie  other  iiillnciices  all  tciiil  to  iclaicj  the  rcjiction  am!  pioirressively 
lii\(er  the  value  of  K.  Ncnative  aiiloeatalysis  oci-nis  when  the  cn/yiiic 
|iiiii|iiccs  pniiincts  whieli  interfere  with  its  activity.  (Jraiiiial  destriic 
lion  of  the  en/ynie  ami  its  union  with  the  proihiets  of  its  activity  will 
manifestly  also  (lecrease  its  power.  There  is  plenty  of  evidence  that 
lioth  of  these  processes  max    occur. 


Reversibility  of  Enzyme  Action 

I'.iit  the  most  important  of  ail  the  causes  of  retardation  of  enzyme 
activity  is  uiidonliledly  n  ri  rslhilil i/  of  iirlinii,  which  is  an  application  of 
the  law  of  nias.'<  action  (  pa^'c  "J.'i  i .  If  we  taki'  the  .saponilieatioii  of  an 
inIci',  the  cipiation   is: 

cii  cii.i'ii  i-oor  III!  otr,  ni  ni  ni  coon  .  (■  II  (Ml. 

(I'thvi  I.mI.m!cI.' I  I  l.utMir  .-ii'i.l  I       ( illi\l  .il.-..!!,,!  1 

I'iic  eipiililii'iiiiii   jioint    is   not    so   mar   the   position   of  complete   hydrol- 

.vsis  as  in  the  case  of  the  inversion   of  saccharose;   in    other   W(n(ls,  the 

•ciidcncy  for  the  liodics  produced  hy  the  hydrolysis  to  reunite  and  form 

tin'  orit;iiial  siilistances  is  ipiite  marked,  so  that  the  reaction  comes  to  an 

I  nil  hefore  ail  the  ethyl  Inityrate  has  licen  decomposed.      l''or  some  time 

licfore  the  ei|iiililirinm  jmint  is  reached,  there  will  have  existed  a  proiires- 

^ivcly  increasinj;  opposition   to  the  lircaUdnwii   of  the  ester,  as  a   eoiise- 

i|iieiice  of  which,  when  en/ynies  are  used  to  accelerate  the  reaction,  the 

Velocity  constant,  as  determined   li.v    the  aliovc  e(piatioii,   will    irradlially 

i.dl  as  the  reacfion  j)roceeds.     ("onveisely,  in  a  mixture  of  eth.vl  alcohol 

mil  hilt  V  lie  acid  tiiere  is  very  slow  synthesis  to  ethyl  iiiitvrate,  and  here 

I'.'aiii  lipase  accelerates  the  in-ocess;   it    induces  a  recoiriii/alile  synthesis 

•itliiii  a  short  time.     lOtliyl  Inityrate  is  usually  em|)loyc(l   for  these  ex- 

i''iiiiients  liecanse,  on  account  of  its  o(hir,  the  ester  is  readily  recoj,'nized. 

!  !ius,  if  the  alcohol  ami  acid  l>c  niix"d  alone,  no  esicc  w  ill  hi'  detectahle, 

^  'it   if  some  lipase  lie  added,  it   will   soon   Ipcconie  so.      Similar  syntiietic 

^I'tioii  of  ii])asc  has  also  heeii  dennnist  rated  I'm-  iikmio-  and  tri-idein. 


i  "*  lin  ^Ic  (11  III  MIc   M     i;\-.|-    III        IIV^Iiil  111,11    \i.    I'Hiii  I  >si:,s 

ll   ^liniiM   III'  i-liiillv    iiiiilcisl 1   llial    |iillf  riit;il\  s|n.  surli   ;is  tlir   liyillc 

L'fll  iiiM,  III  ;iriTlcl;ilii|.,r  ;|  i,.;i,'|jiili  l;kr  lit,,  iiliinc.  i|i.  m.  ii|i|Jllly  ill  lioll 
iliici'liniiN,  SI.  lliMt  till'  |iii>.itiiiii  111'  ri|iiililii  iiiiii  I'i'iiiiiiiiH  liiii-liiiiiucd.  I'lii 
/\  llli'«.  Iiiii.v  .  hiivv  i'\  I'l,  iMii^f  tills  |iii>iliii|i  III  r|i;i|i!.'i'  lii'cillsr  iil'  tlli'ip  liHIii 
illU'   ilitrl'liii'iliiiti'   I'lilillillial  lolls. 

'I  III    /■(<■«/■>(    jiliii^i    1,1   iirliiiii   ri  lift  (III, s  is  iiii,l,iihl,i  III,    ,,iii\,    1,1   III  1,,,^ 
Slum    III   llti    sii„ll,,l,r  firni  I  ^M  \   irlmli   nniir  in   II,,    iniiiiinl  huihi.      A   <ivr,\- 

ilil1ii'iilt>    ill  :iri'i|.tiii._'  Mii'li  a  \ii'u.  Iiuui'mt.  is  llu-  I'jifl   lliiit    tli |iiilil. 

I'i'Hii    |iniiii    111'  all    IimIiuI.v  lii'    nai-limis,    in    tlir   |iifsi.|irc  of  an   cmtss  n 

walir.   is  so   nrar  iiplili'   IimIimI  vsis   lliat    M'Iv    little   s\  iitlii'>is  ran    ]<■ 

|iiissil,li'.     'I'liat    is   ttui'   SI.   Iiiii!j   as   till'   Mil.staiii'c  sy iitlicsi/ci|    is  siiliilil. 

I'll'    il'  i'    i-"   iii';iil.\    iiisi.iiil.l.'   in    wat.T,  or   it'   it    is   inn lialrly   rcinov.' 

Iioiii  till'  sit.'  1.1'  till'  irai-lioii  li\  iJilTiisii.n.  or  in  aii\  otiirr  \\  a>  ,  tlini  it  i 
ol.\  ioiis  thai  it  \\ill  •.'lion  lii'iiiL' s\  ntliisi/fil  liy  tlic  i-cai-lioii.  Tims,  in  tli 
intistiiir   iinitial    I'al    is   liyilioly/i'il    liy    |ianiTi'atii-   lipasi'   into    f;ilty   iiri' 

.•mil  i;l\  irrin.  wliii-li  ,iir  ,il.si.ilii-,|   into  tl |iitliiliuiii.  \(  licic  tliry  ai.'aii 

'•"III''  im.li'i-  Ihr  iiilliiriiiT  i.r  inti-i Ilnl.ir  lipase.     Tliis  l.-itli-r  will  li'inj  t. 

.lei-i'lriate  tlie  sMiliiisis  of  ni'iiti'.'il  fat  I'liiiii  till'  tatt\  ai-ii|  ami  Silyccii 
mitil  the  ei|llililil  illlil  |ioint  of  the  svsteiil  i  la t  ;ieiil  ul.veeriii  ;=!  lielltl,! 
til'       ll.o     is  aLMin  leaeheil;  lull   tliis  point,  .•illlioiinh  it  is  near  llic  iii.'li- 

li'ili'l   "I   fl Illation,   w  ill    |eal|\    ne\  er  lie  re;iehei|    lor  tllc   feasiili   t  lial    11 

iieiilral   f.il.  as  i|iiiekl\    as  it   is  loll I.  will   I me  i|e|iiisilei|   in   ilisoliil, 

'.iiol.nles  in  the  iirotopl.-isni  ;ini|  thus  I.e  reiiio\eil  from  the  ciination.  I 
sil|i|iorl  of  this  \],-\\  it  has  lieeii  fomiil  tli;it  lipase  is  present  in  iiileslii,, 
iinieosa  after  all  traees  of  aillieieiil  panereatie  jiiiee  liaxe  I. ecu  wash, 
.-lu.iy.     I'..\   similar  re;iel  ions  the  fat  of  the  tissues  lueonies  .leeoiiiposeil  : 

f.itty  aeiil  ami   Lil.M-erin  ;inil   passes  mit   of  the  l.h.oil   when  the  coi nil 

'imi  "f  I'al    in   lliis  lliiiil   falls  l.elou    a  <-eilaiii   level.      I'loviileij   one  of  r 

sill'slaiiees  syiilliesi/nl  is  iiisnliilile  or  e;m   in  sonie  other  way  I.e  re \. 

fioiii   the  reaetioii.  il    is   plain  that,  e\en  ti irli  the  eipnliliriiiin   point 

\ei\     ne;ir  to   lh;it    of  eomplele    h.\  i|ro|\sis,    \  el    the    re\ersin,i    wil|    |„.   si 
lii'iint    to  ill.  all   that   is  reipiireil  of  it. 

ii'esults  siieh  as  the  al.o\.'  have   promptnl   man.v    t iielmje  tli.-it    il 

liy   sueh    re\ersifi|i-   ai-tion    that    all    suilhetie    pn.ei  sses   iieelir   in    the    li\i 
i.r>_'atiism.     p.in   ihi.  ilemonst  raMe  s\  nlhi  sis  of  an  ester  must  not   lie  t.'ik- 
as   e\ii|eiiee   tli.it    all    oihrr   s\  nt  hrses   are    e\l)l;iilialile    on    the    sjiiiie    I).!- 
h'or  e\;iliiplr.  we  lia\e  srrn  aliii\e  that   in  the  e;ise  of  (•.•|iie  sllu';ir  the  e.; 
lihriiim  point  in  the  e.pialion  is  si.  nrar  that  of  eimiplete  liyilnil.\  sis.  thai 
me.isiir;il.|.'  aiiioiml  of  eane  sn'j,ir  is  formeil  when  ilexln.se  ami  levnlose  , 
■•lilor.eii     ill    art     ,.n     ,  ,i,  h     ,,iii,r.    .ilel     thai     i-alie    siiojir    does     not     ,-|pp 
wlieii    sneiMse    is    aihleil    to    the    nii\lllle.       If    iiisleail    of   sllcr;|se    We    t. 
another  of  the  sm.i;ir  en/ymes     miiiiel.v,   mall;ise.   w  hieh   .-leeelerates   i) 


n  IIMI A  IS,    (lie    I  \/.\  MIS 


7f» 


■  |i  c'll|ll|l<isitil>ll    III'    lllllltllM'    illtil    t\Ml    lllllll'l'llll'S    III'    ullli'iiM-        llll'i'l'    is,    IlliW 

■  \  iT,  I'S  idclicc  ci!'  sviillifsiN  iis  ii   ri'siill   nl'  the  iiri'dii  iil  inn   i,\'  i\  ri'\  cisilili' 
irjictiiiii.     To  Miiilcrstjiijcl  ilifM'  icsiilis  \M'  iiiiist   ii'iiii'iiilici-  iIkii  iiii|iiiiii'\ 


■NlriiM'   IS   II    MilXtllli'   III    l«ii   sirl  riiisiiiini-,  ilrMuiui 


It'll  u  :iliil  H.  Wlii'll 
!\\ii  liiiilri'iili'N  111'  II  i|i'\l  I'lisi'  riiinli'hsr  ||i:it  iN,  I'iim'  |iii;lli.r  witli  tin' 
lii«.s   III'  il    llliillM-llJl'    III'    Willcl'     lllilltiisi'    iN    rmilli'il.    Iilll     vvill'll    l\MI    lllllll'l'llll'S 

mT  fi  ijiAli'iisi'  I'liiiiji'iisi'  isiiiiiiilliisi'  ^l■•.llll^,  'I'lii'M'  i>  siiiiii'  runt  iii\  rrsv 
.!>  Ill  \\  lirllirr  iinilhisi  is  iriillv  ii'-.|iiinsilili'  Im-  llir  s\  nlliisi>  nf  „  i|i\|iiim' 
niiilri'llli's  111  iiuilliisi',  it  lii'iii'.''  rliiiniril  liy  ^ninr  llnit  tliis  is  ^ii'i'iini|ilisliri| 
li\    jiiHitiii'i'   i'ii/\  nil',    I  iiiiilsiiii .      ir   liiis    wi-ii'    tiiii'    il     wiiiilij    iiniti'iiiillx 

iiiiniiiiizi'  till'  iiii|iiiit.iiii I'  ii'\  risilili'  iirtimi  iis  ;i   r.-n-lnr  in  I'llhil.ir  s\n- 

'III  sis.  Till'  liltrsi  I'viiji'iii-r  iinrs  In  s|iiiv\  ,  Imw  i\  ir,  thul  it  is  liiiiltiisc 
Mill    nut   rllllllsilii'   lli;il    is   li's|iiilisililr    in    ilii'   ali'ivr   i-,isr      i-j'.    I'lMvliss^ 

I'lv  iili'tirc.   Iiiitii   iliiii't    ii'iil    iinliri'i't,    is   iiUn   sirmlily    iii'i'iiiiiiihiliu'.'   tn 

^liiiVi    tli.it    I'lizyilirs    ni;i>    iiri'rli't.llr    I  lir   svnllii'sis   ul'    |ili  itriiis.       As    |iii'i'i'-. 

'i|   iliiri'l   i'\  iili'lii'i'   ur  liiiM':   ili    tlii'   I  I't.iiiliit  inn  iil'  llir  iliiri'st  iv  i'  jirtimi 

■■••    tivjisin,   I'll'..    «liirll    si'ts   ill    iillir    llu'    |i|iiri'ss   li.is  irnlii'   nil    I'lif    ;i    tilili'. 

.'iinl    I 'J  I     llir    irrnlHIIiriiri'llirilt     nl'    .'I    iliursi  i\  i'    |i|iiri'ss    Jl|i|i:i  li'lit  ly    ill     Mil 

I  111!,   il'  I'll'   jilnillli'ts   III'  till'   iliui'slimi    ,'iii'   irliiiiM'il    liy    ilijilxsis   nf   nllii'i' 

Mil  iilis.       As    ililrrt     rviilrliri'     lli;i>     lir    I'ili'il     llu      I'nnil.'lt  iiill     iil'    svntlli'lii' 

|iiiii|ui'ts    wlii'll     |ir|isill     is    iiijili'il     III    I'lilli'iilt  siihiliiiiis    III'    Iif|i|iini', 

.Hill  llii' iliiiiiMiitinn  ill  till'  iniiiilici' III' siiKill  iimli       li's.  .-is  JMili;i'i|  liy  iin'iis 

irniii'iits  III'  cli'i-tiirjil   i'iini|iirti\  ily.   wlii'ii   tiyiisin   is  :iili|ri|   In   tin-   |iriii| 

111-  III'  lr\  jitii'  ili'_'i'stiiin  111'  i-iisi'iniiL'i'ii.     I'r.it^iiiiini-     ;i  sini|ili'  tnriii  nl"  |irii- 

•  111     lijis  jilsn  lii'iii   I'nmnl  til  III'  |iriiilui'ri|   wlii'ii   li\|isin      nlitMiin'il   rinm 

I    iiiiilhisi-      w  ;is  iiilili'il   tn   It    tryi'tii'   iliiTi'st    nl'  tin-   siiiiir    |iriil.'Miiin         'I'lir 

-i'jiiilii';iii.'r    111'    llii'si'    r.-ii'ts    ill    riinnrrtinii    willi    llii'    nirtiilinlisni    nl'    tin" 

ininn   .'liils    will    III'    f\  iili'lit     w  lirll    Wi'    i-nlllr    tn    sliiil\     lliis    siiliji'i'l     i|i,ii_'i' 

','ls  .    'I- 


Specificity  of  Enzyme  Action 

Altliiiii'_'li  ill  :ill  III'  till'  jiliiiM'  I'l'iitni'rs  III'  rn/\  nil'  iii-linii  llirrr  is  nnthin'j 
'  I'liill-iiijii't   till'  \  ii'W    tlnit   tlii'y  ;iii'  i;it:il\  I  ii-  jiiii'iUs,   tlii-ii'   ii'iiniitis  niii' 

■iiliiirity   wliirli   ;it    liisl    siirlil    sri'ins   nninti'ijui't.-ilili'   nn   sin-li    :i   imsis. 

•iNisx\itii  ri'i.MI'il  In  tlii'ir  iil'li'll  Irlll.irUMlili'  spiTitii-ity  nl'iii'linii.  'I'liiis, 
■    \M'    li;i\i'    SITU,    iiniltilsi'    ran    ll\  illnlv /r    lilllllnsr    alnlll'    iwhii'li    is    I'nlii- 

■-"'I    nl     IWii    u  dl'Xtrnsl'     lllllll'l'llll'S  I.     Iillt     lint      isnlua  1 1  nsi'     ;  I'l  nn  piisi'i  1     III' 

i'-\tiiisi' I .      Tills   iiii'Miis   that    iiiiTi'   ililTii'iiii'i'    in    llir   I'linlitiniiil  imi    nf 

'I'llli'S  is  suflii'iclit  to  alti'i'  tlir  inllllriii'i'  nl'  rii/.>  iiii's  nn   tlii'iii,      Sinn' 

■  !:  ilill'i-i'iMii-i's  i-n\\]i\  !ii>t   ititluiiii'f  that  nf  inniiiaiiii'  *'a'al\~'s  \\i-  miis' 

■''^'     'il>'-    1""     iniM.-     in    lln,    :.,l..,r^il.'rv     I..    .1,  r,.  i,:-n  ,|1.-    ,i!.v     v,i!ll.,.i       ..f    ^\....^,:,     «;-.■,    tlv 
'-  ■  1-    I'l'li'l  I.,  a  liy.lr,,|>.w  .uiMiu.'  .,f  .liMiii.  .  iil.il;..-,'  .I'l.l   ul -.     i.r...|ii.,.|   l.\    ill,    |.|.  |..iil:,  ,| 


1 


MO 


|-|IV>I<  III  III  Mil  \l.    Il\.'-I>   UK    l'insil)l.iNI|r\l,    I'IKM  i>hi:h 


)'\|ililill  tin"  riiPsc  III'  I'll'  ililTiii'lM'i'  Tills  liiis  Imtii  iIiuh'  nil  tlii'  liasi^ 
citlli'l-  tlllll  I'M/yilii's  ill'r  I'lillniiU  III'  llinl  lln-  iirlivi*  (i-iltal\tii-l  Ul'iillp  III 
tlic  rlizviiii'  is  iittiirlii'il  III  a  nillniil  iiiiiIitiiIi-.  Ili'l'iirc  a  siilislaiii'i'  i-aii 
III'  ai'ti'il  nil,  il  must  I'liiiiliiiii'  with  lln*  I'lilliiiil,  whii'li  it  ijnfs  liy  tlit>  prm- 

I'MN   m|'  a<lMi>i'|itiiiii    (MT    |ia(rr   li.'ii.      Tliis  I'iiii   1 iir,    Iihwi'mt.   iiiiI\    wIu'i, 

tlii'i'i'  is  a  liai'Miiiti.\  lii'tw  ('I'll  till'  nilsni'liini;  siilistaiiri'  ainl  llir  siilislaii"< 
ailsiirlii'ij.      iiistiiiii'i's  III'  till'  spi'i'ilii  it y  nl'  ai|siir|itiiin   liavf  alri'iiiiy   licrn 

L'iM'll. 

Ill  sii|ipiii-t   lit'  this  xii'w    it    lias  lici'ii    rmiiiil    tliat    nf  llu'  two  iimhitsi^. 
II   ailij    H.    ill    till'   spli'i'll,   mir    is   ailsnlliril    Imt    lint    till'   iitlll'l'    \\  lli'll    a    siilli 

timi  I'liiit.iiniiiL'  tlu'in  is  sliaKi'ii  with  Kii'si'l;(iihi'.  Kui'lhi'i-iiini'i-.  whrn 
siiliitiiiiis  lit'  iii\rri,.>t'  air  shaKi'ii  «illi  rt'i'taiii  iiicif  pnwijirs,  the  in 
Ni'itasi'  is  ailsiii'lii'il  liv  siiitji'  111'  tlii'in  Imt  not  li>  iplhcfs.  In  sti'mii;  slip 
pull  111'  till'  a'lsiirptiiHi  hvpiithi'sis  is  also  tin-  t'art  that  tin-  same  mathr 
niatiral  laws  ;  apply  in  tlir  p"ii'i'ss  iil'  ailsiiiptinn  aii-  iilii\\ril  in  tli' 
I'alin  x»hii'h  csisls  lirtwi'di  tin-  ai-livity  nl'  an  rii/.\  iiic  ami  its  I'lmci'ii 
li'iitioii   ill  till'  siihiiiiiii. 

'I'll   sMiii    np,    thi'ii,   ratal.Nsis   as   i'\|iiliiti'i|    liy   cn/ynics   iiixnlvi's   thii"- 
piiii'i'sscs :  i  1  I  I'lintai't  lii'twri'ii  till'  rii/\iiii'  ami  tln'  siilistrati',  whii-li  will  I'. 

ili-prnilrllt   nil   tlii'ir   rail's  nl'  ililVllsinn  ;    I '_' I    ailsnrptinii   lii'twi'i'll   tlli'lll.   whii  II 

will  ili'priiil  nil  tlii'ir  I'nnli'.'iii'aliniis  let'.  Ihf  Ini-k  anil  ki'v  similri;  an'l 
I'ii  the  r'li'liiii'al  i'halii;r  w  liii'li  itst'll'  pinlialily  taki's  plarr  ill  twn  stiiL'i  - 
In  rniini'i'tinn  with  tin'  thinl  prnri'ss.  it  is  pinhalilr  that  an  initial  I'l'in 
pniinil  nl'  a  ili'liiiili'  I'lii'iiiii'al  natni'r  is  lii't  I'niim'il,  rnllnwi'il  hy  th' 
hyiltiilv  I  ir  111-  nlhi'i-  rhcniii'al  rhailtrr,  al'li'l'  which  llii'  rli/.yiiii'  jjimin 
liii'iimrs    t'l'i'l'. 

It  is  \('iy  siiiiiidranl    in  this  I'ninirrtinn  In  nntf  that   in  their  snliihi! 
itii'S   thi'ie    exists   a    ilistinet    li'lat  iiiliship    lietween    the    I'll'mellts    ailil    tl" 
sniistiati's  nil   whii'h   they   reaet.     Thus,  tr>p>iH   is  very  sniulile  in   \(;i' 
ami  ai'ts  nii  w  aler  snliilili'  prnteiiis:  lipase  is  sniillile  in    I'at   snlveiits. 

Certain  Peculiarities  of  Enzymes 

Nnl  w  ilhstaiiilill!.'    the    \i'iy    strnnir    ease    that    is    niaile    nut    t'nr    the   e,-' 

l\iii'  h>  (inthesiN,  theie  are  eertain  I'aets  which  many  liml  it  dillieuli  ■  ■ 
iiiald'  cnnl'nrm  with  sm-h  a  view.  One  nl'  these  is  that  ilextrnse  ■  i 
Uliileli.'n  three  ilistilii'l  aiiil  sepaiale  types  nl'  ileenmpnsit  inn  aecnpl  .; 
tn  the  en/.\  liie  allnweil  In  act  nil  it.  'i'iiese  ale  alciillnlie  rermcntal 
liiilvrie  aciil  t'ermeiitatinn  ami  iaelic  aeiil  I'ermeiitatinn.  It  is  dil'li'  ' 
In  see  how  simple  catal\tie  act  inn  can  he  I'cspniisilile  I'lir  all  three  resii  ■ 
The  eii/yme  must  lint  niily  initiate  the  ehaii^es  litit  also  ilireet  '■'■  '■' 
cnlllse. 

Annthcr  jieenliarity  is  'hat   when  certain  eii/ymes     e.  j.'.,  renniii,  |     ■ 


« 


KKHMfNTS.   <ii;    IN/.V\IIH 


HI 


,iii,  f|i'.     art-  iniiiMi 


IiiIkI 


ill  I   liiiials.   Ilicv    ciiii'x'  «.|ti'i-i(ic  iiiilii  11:111111  s   III 
i|.|ii',ir  in   llir  III I  <'l   tlir  iiMHMiiiiltil   iiiiiiiial.     TIhih,   uIhii   aiitinimin 


I'liiM   in  ailili'ii   to   milk    it    k'liii 


ih    hiiiili'is  cliiltiiiu  'Ml    •''<■   NiiliHi'ijiH'iil 


M>i<lilii>ii  "if  iTiiiiiii.      It   i-'  iiniliiilili'  lliat    piiwci  liil  antii'ii/.,Miif»  an-  pin 

ilur.'d  ill  lilt-  animal  Imily  U>v  tin-  |iiii|i<im'  nl'  |MotiTliiiif  llii-  linMirn  fr 

,ill!ii'k  li.\  I'li/.MiM'H,  It  io  nil  aii'Kiiiil  .if  llic  iircHi'iiiT  "f  aiilii-ri/ymi's 
tluit  iiitrNliiinl  iiaiiiHitcs  ciiii  cxixl  in  tin-  iiiti'stinr.  ami  llif  imiiiiinitv 
iiolii  ilitfcsli.iii  wliifli  till'  milfiisa  <<(  tlir  '.'astniiiili'stiiial  Irart  ciijnvH. 
Is  iiflirvcil  I.I  lir  (hn-  In  tlif  sniiit'  raiiMv  Itiil  tliiTf  is  .■nn>ii.|i'iiilil.'  ilmilit 
i.'.'iii.lini;  tliis  I'liiiin.  Klrsli  |iiiii<Ti'iitic  .jiliri-  wlini  injicti'l  into  tin- 
riii|ii\  iiilisliiH-  iliircsts  ils  walls.  Wlifii  f"".!  is  |iics.-iil  in  tin-  intcH 
line  il  I'viih'iillv  pi  ivfiils  iliirrsti.pii  ol'  Ihc  walls  1>\  ilivcitinir  tlir  iii/.v 'in- 
1.1  itsi'ir. 

Types  of  Enzyme 

IFavinir  Irarnt'il  simiflliiiiir  aliniil  llir  !;i'm'iiil  iialuir  "t'  i'Iiz.mih'  arlion, 
wv  may  imw  turn  i>iir  allnirniii  li>  rtrlaiii  ilri:iils  l!iat  liavf  a  pnirru-al 
nil,..  ■.<•<•.  Ill  llii'  liisl  plai-f.  willi  ic'.'aiil  I"  mum'nrlaliiii',  in  tin* 
r:\\Vw  ,\iiik  carli  iu'wIn  <lisfi><  ficti  fii/.ymr  icccivi'il  a  mime  uliii-li  «as 
(111(11  i|nili'  iiiap'Miipiialf.  Many  »(  llicsc  names  an-  ri'tainctl,  sm-li  as 
prpsin.    trypsin,    plyaliii,   rte.,    Iml    il    is    m.w    iMislnmaiy    to    iiann-    Ilii" 

iii/.ymc  aiToriliiin  tii  tilt'  siilistai 1  wliii-li  il  ai-ts.     'I'liis  is  ilmic  imIIht 

K\  icplariiiif  tlic  last  pail  iil'  Ilir  name  «>!'  tin"  snlistaiwc  arlcil  on  liy  tin- 
I'Tiiiinatioii  -asc  (for  I'xaiiiplf,  tlir  enzyme  wliieli  iiiveils  maltnse  is  ealleil 
iiiallase).  nr  li\  merely  aiiilinir  ase  In  the  name  "I"  llie  siilistaliee  aeleil 
upon  itliiis.  the  enzyme  uliieli  liy<lioly/es  •rlyeoocii  is  ealleil  ^'lyeou'eiiaseV 

MosI   of  the  en/.ymes  in    the   animal    lu.ily  aei-eleiale   hydiolvtie    jiroe- 

esses  ami   are   elassitie-l   ai tlini:   to   the   chemieal    Mature   of   ihe    siili 

strate  on  wliieli  they    work.     Thus,   we  have: 

1.  The  ainijldiitu  aeeeleratini;  the  hvilrolysis  of  iiojys.ieeliaiiiles,  i'.  jr.. 
pl\aliii  liii  siilivai.  aiiiylo))siii  1  in  paiierealie  jiiieei,  yiv  eoi;eiiase  (in 
lixeii.  iliastase  ( in   mall  >. 

■J.  The  iiirirliisis     aeeeleratiii!.'  liyilrol>sis  of  disaeili.irides,  e. -_'.,  malt 
.iM'.  lailase  ami  suer;ise  I  in  sueeils  eiiteiieiis  1, 

;'..  The    i>niliiiiii.iis     aeeeleralini,'    hvilrolysis    of    proteins,    e.  jr.,    ]ieitsiii 
111    L'astiie  .jiiieeK   trypsin    (in    pain-reatie   .jiiieei,   ere|isiii,    inlrai'elliilar 
i'li.teiiiases. 

t.  The  /i/»((.v(.v     aeeeleratiiij:  disruiit ion   of  neutral    fats,   e.  jr.,   steansin 

■I;   pjiiii-reatie  jnii-ei,   intraeellular  lijiases.. 

"1.  Anjiiiast      H'-eeleralinjr    hydrolysis    of    aririniii    into    iiie.i    jiiid    or- 
Ml  hill,  (intraeellular)- 


juj^na 


.S'> 


I'llVSIClM  IIIMICAI.    1'.\S1S    (ir    I'llVSKll.OdlCAI.    I'lilHKSSI'S 


(>.  I'nd^i  jiri-i'IiTJitiny  liy.liiilv  sis  nl'  ni'ca  tii  .■iiriiiiuiiiiiiii  i-iulMiii;iti' 
(in  iiiniiy  niii'iiioiirJinisiiis  ami  in  tin'  soy  licani. 

7.   HI 'I'lxiiliisi      ciinvrrlinir  ulynxals   into   lai-tic  arid    (pa!.'"'  <ili(>). 

(»tii('r  cn/ynn's  ni-i'dcratc  oxiilativc  iiioccsscs  ami  arf  oallod  orlihixis 
and  }ti  ri>.iiihisi  >t.  (Mln'is  Iniiiir  aliouf  tin'  displaci'iiwiit  of  an  amino 
t'l-onp  liy  liydrosyl  iih sdiiiidnsi s) .  (MImm-s  cause  coagulation  (rodfiiihi- 
tin  fi  riiii  iih'< .  c.  tr..  tliiond>in,  rcnnin.  <  Mn'  ol'  tlic  cn/yrncs  pi'cscnt  in 
snccus  cnlciicns  acts  liy  convcrlini.'  the  /Nnniycn  ( ti\  psimiizcn  )  into  tin 
cii/ynii'    I  t  ryjisiiO  . 

Enzjnne  Preparations 

So  I'ai-  it  lias  liccn  inipossililc  to  pi-cpai'c  cn/.ymcs  in  a  pnrc  stato  al 
tlioutrli.  Iicin'.'  colloidal  in  natnic,  tlicy  aic  I'cailily  jn-ccipitatcd  of  ad 
soi-Im-,!   aloiii;'  with   other  colloids. 

Since  most  ciizynn's  exist  in  cells,  it  is  nccessai'y  to  hreak  up  tln^  cell- 
in  order  to  isolati'  the  enzyme.  This  is  done  in  various  ways.  Ry  on^ 
iiH'thod  the  cells  are  L'l'onn"!  in  a  nmrtar  with  line  sand.  tluMi  made  inl" 
a    paste    with   inl'nsorial   cal'th    i  l\ieseli.'ulir>.   thi'    paste   elicloseil   in   stnu' 

canvas  and   pla I   under  an   hyilrauli<'   pi-ess  at   aliout   otH)  atnios]ihere^ 

in-essure:   a    clear    lluid   s<'parates   and    this   contains    tlu'    enzymes.      Am 
othei'  \\a,\'  is  to   freeze  tin'  tissue  with  licpiid  air  and   <rrind  it   in  a  slei  I 
nioi-tar  liy  nn'ans  of  a  iinichine.     Still  another  ami  less  e\i>ensive  method 
;ind   one  \\hich  \\  e  haxc   found    nmst   useful    foi'  (M-<;ans  and   tissues,  coii 
sist:.  in   icdnciiin   the  tissue  to  a  pulp  and,  after  sii'vintr  it   to  iret  fid  i'' 
'■(nnu'ctivc   tisstn'.    etc,   sprcadiuir   the   ]>ulp    on    L'lass   ])Iates  and    df.vin'. 
in  a  sliuditly  warincfl,  dry  ;iif  current.     The  scales  of  dried  material  ar 
then   iri'ound  in   a    paint   mill    with   toluene,   .mil  the  rcsnltinj;  sus)>('nsi.. 
filtered:  the  powder  which  renn'uns  on  tin"  filter,  after  thorouy;])  washiir 
with  toluene,  is  diied  anc]  kept    for  future  use.     The  tolui'Ue  removes  a' 
the   f;itty  snhstance-.,  mi  th;it    when  shaken   with  water,  etc.,  the  enz.\  ni' 
di-.sol\  ('. 

Conditions  for  Enzymic  Activity 

Reactions  lii'oU'_'lit  alioul  lp.\  intracellular  cnz.v  nics  are  very  readi 
iidiiliifeil  wiH'n  there  cmnes  to  he  a  certain  ac<'nmulation  of  their  |iio 
iict^  of  action.  Thus,  yeast  ceases  to  I'ernH'nt  su^ar  wlu'ii  tlic  alcnh 
has  aceuiiiulaled  to  a  cerlain  i)ercenta'ue.  This  aclion  is  partially  d 
to  a  toxic  action  of  the  alcohol  on  the  cell,  \\hich  paialyzes  its  ]iow  er 
alisorliiiiL'  the  snlistahcc  to  lie  acted  on  hy  the  intracellular  enzyme.  I 
these  pi-odnels  lie  not  in  some  wa\'  fenioved.  they  will  ultimateh  I. 
the  cell  :ind  stop  the  feiirn'ntal ion.  We  ha\e  seen  alioNc  how  the  ;: 
ciiniulation  of  prodiicts  may   inti'ilVrc  with   the  activities  of  enzymes 


n  mil  NTS,  (Hi  iN/vMr.s 


s:t 


.i|li(  r  \\ii\s  in  wliicli  tlic  cii/.*  mh'  doi's  imt  MitVcr  ili'stnicti<p|i.  iis  is  slnwii 
li\-   till'    liicl    tiiiit    it    I'fMiiiK      i1s  (iiiiriiiiil    iiftixitics   mi    iciiioval    «>i'  tlic 

pl-dllui'tS. 

Hn/yincs,  Imtli  iiilrju-cllulMr  iiml  est  i:i<'ilinl;ir.  mi-o  very  sensitive  tn- 
uiiriis  tlic  iniirsiiinic  rip|iii>.>sition  <it'  tin-  iiii'dinni  in  wliidi  llicy  iirc  .'ict- 
iiiiT.  Vur  tlic  iiitriicclhiljir  cii/..\iiics  this  is  wlint  \vc  slumlil  c\i)cct  when 
we  ItcMi'  in  iiiind  tlic  i>nit'iiilii<l  iiillileiice  of  ini>f";iini<'  sjilts  on  llic  lieart 
l.ciit  and  on  cell  "ji-owtli  and  division.  Tliis  inllnencc  of  salts  ami  of 
ivaetioii  (acidity,  etc.  i  on  llic  life  of  tiie  cell  is  so  iiroiioiineed  as  to  lead 
sniiic  oliservcrs  to  liclievc  tliat  alniiH'iiial  cell  iiinltiiilication  in  tlic  lioily, 
,is  in  tlie  ease  of  liinior  i'ornialioii.  is  due  to  clianu'cs  in  the  inorjianic 
i-iiiii|)osilioii  of  the  tissue  tliiids.  Kxt racelhiiar  eii/ynies  are  also  very 
siiseeptiiile  to  the  inllileiicc  of  inoruaiiie  salts  Imt  niofe  cs|>eeially  so 
towards  tiic  icaetion  of  the  solulioii.  In  tcniis  of  inodcrii  clieiiiistry 
we  may  say  that  the  eoiiceiiti  alioii  of  If-  and  Oil'  ions  has  a  inofoiiml 

iiillucn II   the  activities  of  cn/\iMcs.     .Most   of  the  eii/.Miies  of  the  ail- 

iiiiai  \'<i\\  ))erforiii  their  action  inniiialix  in  the  |iresciicc  id'  a  sliudit  cx- 
less  of  oil'  ions,  tliat   is.   in   faiiitlx-  alkaline  reaetioii.     Indeed   the  only 

.  \ce|iiion  of  iiii|iorl:ii to  this  is  the  iicpsiii  of  i:as1i'ie  .juice,  which  mn-- 

i.i.iUy  acts  ill  .-111  acid  nicdinm.  .\ii  excess  of  cither  Oil'  or  II-  ions 
iidiiliits  the  activity  of  the  enzyme  and  usually  destroys  it  peiiiiaiiciitly. 
Tlic  aetivilies  of  ciizviiics  are  also  iiillnenccd  l>y  li','ht.  many  (d'  Ihi'tii 
III  inu'  destroyed  liy  siinliulit :  cclU  siieli   as  micioorizanivms  arc  siiiiilarl,\' 

llfl'i-cted. 

I'.cfin-c  hciiisi  secreted  the  diilotiM"  cii/.yiiics  exist  in  the  cells  which 
iir.MJiicc  them  as  inai-tivc  iireeiirsoi-s  c,-illed  :iiiii<i!i(  nx.  The  i;raiiiilcs  seen 
ill  rcstiii-i  udand  cells  arc  (d'  this  nature.     The  activation  of  the  zyiiioircn, 

■  •V  its  conversion  into  the  enzyme,  occurs  after  it  has  left  the  cell,  and 
'Ills  has  hci'ii  coii-.iilcred  as  another  safc'.;iiard  to  dicestion  of  the  cell, 
-".iiii'lim-'s  the  activation  does  not  i iir  until  the  z,\iiii'Lren  has  travelled 

line  disl.-inee   jiloim   the   liland   dllcl,   as    in    the   ease   id"   the    proteolytic 
I, /Ml f  panercatic  .juii-e.     Till  it   reaches  the  intestine,  this  exists  as 

■  \psiiio!ien   Itlii'  zymou'cni.  hut   it   is  here  acted  on  1),\-  iiiiother  enz.vmc- 
:.i    liody  iirodiiecd  l>y  tlie  intestinal  epithelium  and  cjilled  cnterokinasc. 


rilYStCOCUKJflCATi   KKI-  KI.'KNCKS 

(Moiioi^n-ai)hs  ;ind  (>riuinal    I'apcrsi 

■.:Ns,   W.   M.:      IMiirii.l.'H  (if  (i.iTiiTal   I'li.vsiMl.iuy,    l..iii^in:,ii-<,   (inm    \    I'e.,    I',il'. 
ill.,  .1,  C:      I'livsii-MJ   Ciiriinsti-v,    lis    lii.-oin^   mi    l'.i.ili't;,v    iiihI    \I  .   li.  in.-,    AmhiIiI 
i-l.  •:.  Il'll, 

'   11  miull,  .1,   .•^.:       l'liV-ii-;ii    I   |i,i|nisliv    m     X'itfi    I'lu- mm,    I'i  in.  i  Imi     I    iii\.i-il,> 

I'l—s,    PUT. 
I'll"-,  i:.  II.:      IMti.1|i|,.s  (,r   lluneiii    I'liv >i.il..-y.  r,|.  :.',    IIH:.,    I..i:i   :in.|    K.liiv.i-. 


84 


I'llVSIriirlllMHAI.    IJASIS   (>P    l'IIYSIlll.(>(lll'AI,    l'H(H'i:ssi:.s 


I 


.1.  <1 


I'ltirc' 


KalilcrilMry.    I,.:      .Lmi.    I'livsiral   (liciii..    MmmI.   \.    111. 
•■I.'i'i.l,  K.,  \\  i\  iriciiiili:     .Iniir.  riivsicil..  Is'.is,  s\ii,  hi. 
;\\il-iiii,  'I'.    .M.:      Am.  .Imir.  I'liv^inl,    IIHI.'.,   \;ii,    l.'iii. 
■Il.il.hirii-.  .1.  S.,  :iii4    I'liistliv.  .1.   (I.:      .l,,iir.    I'livsii.l.,    l!i|i;,   I    \1W:    IViiMtlcv 

II. i. I..  |..  .;ii|. 
■rliiik,  W.  M.,  Mini   l.nlis  II.  .\.:     .Idiii.  Ii:iilrr  i(.ln;;v.   I!il7,  ii.   I  mi. I    luil. 
1'  II. 11. 1. IS. 111.  I..  .1.:     Till'  Kx.iTlii.ii  i.r  All. I   ill   llriiltli  :iii.l   Disi'ii^c.  llurM'v   I..-' 

.1.  H.  l.i|.|.iii.'.itl  <'i...  Il'l.'i.  X.  l:'.J. 
iilliii.iiisi.ii,  1,.  .1.:     TIm-  I'itiirvs  .,r  till'  Kii\ir..iiiii.'iit.  Mnciiiilliiii,  \,   V.,   l!H:! 
i-\:iii  SIvk.-,  I).  I>.:     .1(1111.   Hi.il.  Cli.iii.,    I!il7.  xvx.  L'sii.  .;I7. 

1  i.iv.v,  i;.  I,.,  :iii.i  K'l.HiiiiTc,  I..  (1.:    .\,,ii.  iiii.  \ic.i.,  iini;,  xvii,  r,.'.-!. 

iMiill.ii.  (i.   i:.:     .1.1111.   lii.il.  Clniii.,   l:i!7,   >\\,  :w.\. 

1    I'.iliiMi-.  \V.   \V..  nil. I   Ih.ii.l.TM.ii,   I,.  .1.:      Ai.li     Int.   Mc.l.,   IIU.;.  \ii.   1.-,.!. 

"S.llar.ls.    A.    \\ .:      Tlic    l'iiii.i|.|.  s  i.f   A.i.li.si.s   nii.l    I'iini.-il    .M(llii..|-.    I'l.i-    Its   Stii.lv 

ll:tr\;ir.l   l'iii\  cisit.v   j'rrss.  ( ';iiiiltri.lj;i'.   Ii»l7. 
i"l.li.v.l,  I".  II.:      rii\  :iti'  <'<iiiiiiiiiiiii'.'iti..ii. 
i-M:ir;illiifii.   .\.    U:      Siiif.'ii'f  Tiii^ini.   .'iiiil    \il;il    I'li.'iii.iiii'iia.      riii\ri>.it.v    ..r  T 

.^luiliis.  N...  s,  IIHJ:  nisii  Kru.l.iiis*..  .I.r  l'li.vsi.ili.;;ii..   I!i||,  ii,  ."ills.' 
i''li;i.vli-s,   \V.    M.:      Kii/vnii.-    Aitii.ii.  r.l.  J.      Miiii..:;im|iIi«   in    Uiiii-lii'iiiisl .  v.    I.nii 

(irrili    i.    Cii. 


iniiit.. 
111:111^. 


PART  II 
TIIK  liLOOl)  AND  TlIK  LYMPH 


CIIAITKU  X 

r.Looi):   ITS  (IKNKUAI.   1>I{(  M'KKTIKS 
r.Y  H.  C.  1'i:a»(1,  It.A.,  M.I). 

Till'  lildoil.  l)('iiiu  tlic  ciiri'icr  of  tlic  niiliitivc  iiiiil  wiistc  siil(st;in('(>s  of 
tli<>  IiikIv  's  iMctiiliiilisiu,  must  iit  line  tiiiit'  or  aMntliiT  cdiit.'iin  nil  tlu>  iiia- 
ti'i'iiils  w  liicli  (M)iii|>i)S('  till'  tissues  in  aiMitinn  In  tiiiisc  uliicli  arc  jn'culiiir 
t(i  tilt'  liloitd  itsclr.  It  is  ii  very  imhhiiIcx  tlui<l,  ami  all  dl'  its  ciiiistituciits 
lire  iii)t  fully  kiiiiuii.  Stiurturally  ii  is  (•(iiii|'"  I  <i\'  watci-  in  wliicli  arc 
dissolved  various  irases  and  orfraiiic  and  inn  nie  liodies,  tlie  corpuscles 
;ni(l  platelets. 

THE  QUANTITY  OF  BLOOD  IN  THE  BODY 

The   most    accurate    Method    ol"   deterniinini:    the    volume    of    iihiod    in 

the  hody  is  liy  lilccdinsr  and  sulise(|ueiitly  washiiii.'  iput    the   lilood   from 

ilic  vessels  and  then  estiiiiatinir  the  amount   id"  hemo'_doliin   in   the  totai 

lliiid   (Welclier's   method  i.     Tins   mctliod   employed    in    the   case   of  two 

criiiiiiials  mIio   liail    liccn   deeaiiitatcd    irave   the    wciylil    of   the   Mood    as 

7.7  and  7.1!  jx-r  cent   of  the  hody  \veii:ht.     I'.loodlcss  methods   for  ileter- 

niiiiiui;  the  total   volume  of  Mood  are  liased  ui>on   the  iirincijile  of  add 

iii'_'  ;i  dctinite  ([uantity  of  a  known  sulistancc  to  'he  circniation  and  tlu'n 

1  stiiiiatiuy:  its  concent lation   in   ,i  saTiiple  of  lilood  withdrawn    from  the 

'"Illy  shortly  afterward.     If  the  sidistancc  can  not   leave  the  lilood  vessels 

;iiiil  dues  not  cause  tluiil  to  lie  withdrawn  from  the  tissues,  the  total  ipnintity 

•f  Mood  in  the  liody  can  lie  calculated   from   the  concentration   of  the 

iii.H'ctcd  sulistancc  in   the  Mood.     The   most   accurate   methods  liascd   on 

'his  principle  are  llaldanc  and  Smith's,  in  which  cailioii  monoxide  lms 

>  inhaled  in  a  '^iven  !;nniunt  and  the  cailmn  monoxide  lienioirloliin  snli- 

-ii|uently   determined    colorimetricallv  ;    and    Keith,    IJowiitrce    and   (ler- 

iLrhty's,  wlucli  employs   vital    red.   a   dy(>  of   low   dilTusiliilily.     The   d\  c 

"iiiaiiis    loni^   I'tiounh    in    the    body    to    lie    ihoroiiiihty    miMi!    \»ilti    lln- 

i'loivil,  and  its  concentration  in  the  plasma  is  d«'termiiied  coloiimetricall> 

S.'i 


8G 


■nii:  r.i.dun  ash   imi;  i.ymi'II 


liv  ciiiiipiiiiiiLr  w  itii  II  Miitiililc  si  ,11 II  1,1 1(1  III  i\t  lire  n|'  dye  iiml  miuiii.  'riicsc 
inclliiMls  vri\i'  tlir  |ii|;il  ;iiiiciiiiil  iit'  IiIikmI  in  till'  IiimIv  jis  ri'uiii  'i  to  S.S  jut 
(•(■III  ol' its  «ci'jlit.  Mrck  liiis  ici-ciitly  ili'v  I'lnpid  ;i  nii'llinil  in  wliicli  v;mii 
jic'U'iii  is  usi'il.     Alter  iiii\iiijr  with  tlif  MihmI,  tlir  I'diicciitiiitiuii  of  this 

sIllislMlli'C  is  ilrlrllilillfil  riniii  the  i-.-ilcimil  ciiMlclll.  Ili'illi;  rollniii,  liDlir 
III'  tlir   iiliiii    li';i\cs  till'   liJiMiil   scssi'ls. 

'I'lli'    llrwiT    liirlliiiils    1m\('    sliiiwil    thill     till'    MiIlllilC    111'    till     cil'i'llliltili'-' 
lliiiil  is  iiijiiiiliiiiicil  riiii-ly  ciiiisljiiil   in  spili'  nl'  inlliii'iii'i'-i  ti'i.dim:  to  ;ilti'i 

il.      Till'  lioilv   :i( iiip!isli(s   this   liy   ilniwiiiy    ii|>iiii    llic    reserve   lliiiil    iii 

the  lis,sll"S  Mini  li.\  \ai\iMu-  ll|e  r.ili'  ul'  w.i'er  eNei'i  iii.ii,  |i|irtiellhirl\ 
tlu'iniiiii  the  kiiliieys.  'N'ejirs  ;i'_''(i  Ihe  duel  line  of  .•in  iiicfeascil  iiiiiDniit  nl' 
iiliKid  ill  Ihe  lindy  i  plel  liur,i  i  y:,'i\e  rise  In  the  theijipriir'e  use  nl'  lileediri'j 
Kspeeiiilly  vviis  this  tliii\ii;lit  In  he  useful  in  cniidit  imis  whieh  we  imw 
ree(py:nizi'  ;is  ehriinie  li\  perleiisiini,  jiiid  whieh  show  iiii  inere.'ise  in  lilnml 
\ipluiiie.  Indeed  \;iriiitiiin  in  lilnnd  Miliiiiie  is  iml  eniiniiipii,  jillhoiiLili 
])lelliiir;i  iiiiiy  oeeiir  in  pnlyeyllieniiii,  ehlnrusis,  mid  ;ii.r  s.  iiiid  theii' 
liiiiy  he  ,'1  teiii]iin  iiry  redilet  imi  in  Ihe  Mlliniint  id'  liliiod  in  dis(>;ises  in 
which  there  is  :i  ^reiit  di'pli'linii  'il'  w;iler.  jis  in  AsiMlie  etmler.-i,  :ind  I'nl 
liiwiii-;  very  sexcie  lienitirrh;iire. 

Wliile  Ihe  liilal  ipijiiitily  nl'  the  lilond  in  tli'  Innly  dues  nut  viiry  irre;ill.\  , 
the  enneelllriltinll  iiI'  its  \,iliiills  emist  illlelils  is  slllijeet  ti)  ilisliliet  eluilliii 
The  Viiluiile  pereeiitjiyes  of  the  eiirpllseles  iilid  the  lil.lsliia  call  he  ilppinx 
iiii.'itely  di'leriiiiiied  1p\'  alhiwiii'^  ii\ai,ili'il  hluud  In  sedinient  nr  liy  eei 
I  ril'imiii','  in  a  '-'iailu;ited  evlinder  hy  the  use  nf  the  heiiialnerit.  Sml 
nielhiids  are  nut  \ery  reliahle.  Iiiit  iii,i.\'  yield  siiiiie  ini])(irtaiit  in rnrinal inn 
Xmiliallx    4.'i   tu   ")(!   per  eelll    nl'  tile   \iiluilie  n|'  liliiod    is  en|ii|)iised   id"  em 

iniseles.      It    xjirles   iiinre   nr   less  direelly    with    the    liniiilier   nf   red    hi ' 

eells. 


THE  WATER  CONTENT  OP  THE  BLOOD 

Sinee   the   Mnnd    pljisiiia   is  essenlially   a   wiilery   snliilioii,   smiie   idea   n 

its    water    ennleiil    e;iii    I hlained    hy    a    deterniinal  imi    nl'    the    speeili. 

irravity.      'I'lie    nmsl    ;ieeur,'lle    iiielhnd    I'nr    aeenlllplishinn:    this    is    In   delt 
lllilie    directly     the    wei'^lit     nt'    a    'li\en     Mihlliie    nf    liliinil    and    cmiipa'        r 
with    the   wei'^ht    i<\'  the   sanie    Miliiliie   nj'    water.      Since    tiiis    iiii'tlind    r^ 
ipiiri"    a    ralher  lar'_'e  animint    it\'   hlnnd.   iinlireel    tiiellrnds   iisiin,'  small' 

aillnlinls    have     hecn    de\  ised.       <  ll f    these     I  Ihininierschljljj's )     uses 

snllllinll  nl'  chlnrnfmin  ;iiii|  lieli/nl  n]'  ;i  speeilie  gravity  of  aliniil  1. •!■''' 
ill  which  ,'i  drnp  nf  Mnud  is  suspended  hy  delixcriiiir  il  caiitimisl,-  fiin 
a  pipette  lielil  at  li'jlit  ailLdes  iie.ar  its  tip.  If  the  drnp  sinks,  ehlnrnfnl; 
is  added;   if  il    rises,  lieii/.nl    is  added  until   the  drnp  renuiiiis  suspend' 


r.i.ddii:  ITS  (iiAi'K.M,  i'i!(>F'r.uTii;s 


.S7 


riic  s|i.'cilic  -iiavily  t>\'  flic  Ix'ii/.ilrlilniol'oriii  inixliiio  is  tlu'ii  (IctiTiiiimd, 
aiul  (his  Viiliic  is  supixis.'.l  lo  v'ivc  llif  spci-itic  iri'.ivity  of  llif  lilocxl. 

Tlu'  spccidi'  •iinvitv  of  tlic  iilnod  <lct.riiiiiicil  in  this  wiiy  Viirifs  lic- 
luiM'ii  1.(140  mill  l.Oti.').  It  is  soiiicwliJit  less  .iftcr  f;itiiiy;  ;iiiil  inciciis.'s 
filter  csncisi':  it  is  sii<;iilly  lourr  liiiiiiii.'  tin-  iliiy  lliaii  at  iii'.'lit,  ami 
ill,,  variation  in  iii.liv  idnals  is  .■oiisiilfialili-.  The  i'lian-i<s  wliirli  oi-cii;' 
ill  the  spi'citii'  !,M-a\ity  of  llic  liloocl  in  discas.-  an-  cliiclly  ililc  to  vaiialioii 
ill  the  pfrcciitairi'  of  pn.'.in,  siiit-c  tlic  salt  coiili'iit  of  tin-  hlond  is  ri'la- 
tivrly  (InciI.  It  is  only  wlicn  u'icat  cliaiii.'i's  oc.-iir  in  tiic  iMincfiit  ration 
,,r  tlic  iioncolloi.lal  salts  that   tlicy  niaiUclly  alTcct   tlic  spccilic  niavity. 

I'l-oin  !M)  to  IfJ  jtcr  cent  of  tlic  plasma  anil  fropi  Mtli  to  (iS.7  per  ciMit  of 

tlie  cnrpiisclcs  consist  of  water.    Of  tlic  whole  hi I.  fr<iiii  tlO  to  7(1  per  cent 

hv  volmiie  or  ahoiit  .V>  per  eeiit  li\  weight  consists  of  plasma:  am!  from 
4t)  to  ;!0  per  cent  hy  vohiiue  or  4,'>  per  cent  hy  wei'.'lit  consists  of  cor- 
puscles. 

THE  PROTEINS  OP  THE  BLOOD 

The  plasma  ohtaineil  liy  ccntrifiijiiii';  the  hhio.l  reiidi'Vc.l  noncoa'.'nla- 
Me  hy  oxalates,  hirudin  or  other  means  (see  iiau'c  !•!•),  contains  .">  to  S 
p.r  cent  of  ooaj-'ulalile  proteins.  These  proteins  arc  serum  allmmin. 
serum  -.'h.hnlin,  and  filirino<;eii.  They  can  lie  se|)arale<l  from  ea.h  other 
l.y  the  use  of  acids  and  neutral  salts.  Their  proportion  varies  under  dif- 
ic  lent   conditions,  hut   is  approximately  as   follows: 


Viliriiiii);rii     .  .  .  . 
Si'Tiini    ;;l"lintiii 
Soniin  iilliuTiiin   . 


a.l.")  <i.ii'~;. 

. .  :'..sr; 


The  aiiiouni  of  lihrinoircii  is  suhject  to  the  ..'rcalest  varialioii  (Mathews). 


"1 


Fibrinogen 

The  least  soluhlc  Of  the  hlooil  proteins  is  (ihriiioncn.  The  plasma  is 
hiiost  freed  of  it  hy  half  saturation  with  sodium  chloride,  or  with  a 
Miall  amount  of  acetic  acid.  It  is  preciititate<l  as  lihrin  in  tli.'  process 
■I  Mood  coagulation  (see  pa-v  !)!>>.  and  is  estimated  hy  wci-hin!-'  the 
:iiioiiiit  of  lihrin  which  i'   produces. 

Serum  Globulin  and  Serum  Albumin 

i.lnhiiliiis  arc  oriiiiiarily  dcrmcd  as  hciny-  iiisoliil.lc  in  «listilled  water. 
il   alhimiiiis  as  hein^'  soluhlc.      It    is.   however,   impossiiile   to  separate 
■  Tuiii  ulohuliii  and  alhamin  salisfaci.,i  ily  in  this  m;iiiiicr.     Tlu-   do'-i! 
II  ohiiiiiicd  hv  diaKsis  can  lie  returned  to  solution  hy  the  ad'litioii  ol 


S8 


nil;  iii.oiih  .\\i>  Tin:  lymph 


ii  suitiililc  iiriiniMl  of  wiitcr.  wliicli  iii.-iUcs  tlic  salt  iKllicroiit  t(»  tlic  pit' 
(■i|<itiil('  M  \.('<'ik  siiliiic  soliii  ion.  In  iitMitral  or  iii'i<l  solutions  it  is  co!)'; 
iilali'il  li>  liral  at  almut  7.'>  ('.  Miit  it  docs  not  art  as  an  iniliviiliiai  i>ro 
Icin.  since  a  porlioii  of  it  is  piccipilalcd  l)y  dialysis  or  l>\  carlioii  diox 
idc.  I'idlialily  sciiiiii  ulolnilin  really  consists  of  two  or  inorc  proteins 
Tlic  sciuiii  aliiniiiin  rciiiainiii^;  in  solution  after  saturation  with  am 
irioiiiniM  snipliate  likewise  doi's  not  represent  a  clicinical  entity.  It  is 
liossilile  |p\  eai'efuliy  lieatinj;  tlic  solution  of  seruiii  alliniiiiii  to  distin 
liiiisli  thiec  separate  coairulatioii  lenipcratlires.  'I'liis  fact  lias  liecii  in 
ti  i|ii(led  as  iiieaniiiu'  that    the  serimi  alliuniili  consists  of  at    least    three 

cluM'l\      lei.ilecj     proteins. 

since    lln    ri friiitiri    null  I-   iif   till    hliiiiil   depeiids    primarily    upon    tic 

a ml   of  proleiii  piescMl.  it   has  lieeii  taken  as  a  means  of  deteriiiiniMU' 

\ariations  in  the  concent  i  ,iii f  the  proteins.      It   has  lieeii   l'<uind  thai 

the  concent  ration  of  ihc  Mood  pioteins  varies  smiicwhat:  during;  ex 
eriise  il  is  increased  prohaMx  liecailse  of  tin'  takiiii;  n|)  of  water  \<\ 
the  tissue's,  and  diiriiii:  profuse  lileediiiLr  it  is  diminished  Itecailsi 
larue  amounts  of  lliiid  arc  lieiiiiz  added  to  the  lilood  from  the  l,\iiipli. 
which  is  rclati\el\  pool'  in  proteins.  The  ini;estioii  of  considcrahli' 
amounts  of  salts  has  liccii  found  to  reduce  the  coiieeiitration  of  tiie  Iilood 
proteins  for  a  sluut  time.  In  pafholoy:ic  conditions,  as  in  diahetes.  when 
lapid  ehiiiiiic-.  in  the  liody  weiirlit  due  to  .dteratioiis  in  the  diet  are  oc 
curriiiy:.  chanires  in  the  lluid  coiiteiit  of  the  lilood  are  often  oliserved 
Likewise  in  edema  <'aused  liy  faulty  renal  function,  there  may  lie  a  ic 
tciiti(Ui  of  lluid  in  the  Mood  liefore  there  is  any  indication  of  edema.     Tin 

hydremic  i-oiKJiti if  the  Mood  can  therefore  lie  considered  as  a  Useful 

diaL'nostic  aid    in  determining:  the  water  metaliolisiii. 

The  relati\ ncentration  of  the  proteins  of  the  Mood  is  also  of  soin. 

interest,  especially  since  in  some  diseases  a  coiisideraMi'  amount  oi 
Mood  piiitciii  is  lost.  |',y  refractrometri<-  methods  it  is  jiossiMc  to  sep 
aiate  the  'jloluilin  ainl  alliumin  fractious.  Normally  the  total  ])roteiii-- 
lanue  lictv\ccn  ii.7  and  S.T  per  cent,  of  whiidi  the  alliiimins  lie  lietxvceh 
4.ll."i  and  77  per  cent,  and  the  ^lolmliiis  lietweiui  I  and  "2.54  per  cent.  In 
sonic  diseases,  as  in  chidiiie  nephritis,  pneumonia,  and  syphilis,  tii' 
ti.tiii    proteins    of   the    Mood    ;irc    dceri'ascd    ami    the    relative    amount    o! 

serum  'jloliulin  is  iiieieased     (In  tl tliei'  hand,  in  man>-  mild  infei'tioii^ 

and  chronic  septic  conditions  tic  uioluilin  fraction  iiia.\-  he  increase. i 
\vitli    I Iiaii'j uiriiii:    in    the   total    protein   content.' 

•  Ilir  kliowled'je  of  Ihe  iir'uint  ilinl  llii  fiiiirliiiii  of  tin  hlnml  firiili  i lis  is 
c|iiiti'  uiisalisl'aclor.\  l'i'i'\ious  to  tin'  discover\-  of  amino  ai'iils.  tii' 
Imililiii'.:  stones  of  the  proteins,  in  the  liloml  it  was  tliousrht  that  li' 
nil  I'oueiiolls    iintlielits    were   i'oli\erlei|    siuiiehow     into    Mood    proteins   dm 


iii.tMiri:  ITS  (ii;ni;hai.  n(iiiM;i{'rii;s 


89 


iiiu'  or  iinirinliiitfly  fiillouiiij;  tlicir  iilisorjitiim  fi'inii  tlic  iiliiiiciitai'.v 
.•.iiial,  !1Im1  tliiif  tlu'  fissnt'  ci-lls  >\t'f('  iionrislicd  i'l'nin  lliis  ciiiiiinnn  pro- 
tein. It  is  tiiiw  kimwn  fluif  the  )iiiiiiiii  acids  ai'i-  luit  iiiiiiiciliatt'ly  svii- 
llictizcil  inlii  1)1(111(1  ))r(itciii«  al'tci'  their  alisorptioii  fidiii  the  div'cstivc 
s\stcm.  Tlic  lil(i(i(l  priitciiis  arc  i-adicaliv  difl'crciit  I'ldiii  the  tissue  jX'o- 
ti'iiis.  Substances  which  retard  (ir  aecelei'atc  nitioiren  nictaholisin  do 
imt  alter  the  relatiduship  existinir  lietwecn  the  |iiiitein  licidies  of  the 
lildiid.  This  fact  indicates  that  the  scrum  prulcins  have  a  function  (juite 
iii(lcl>eiident  of  the  nitro(ienous  nietaholisni  of  the  iiody.  They  uu- 
.liinlitedly  maintain  the  viscosity  of  the  Mooil  and  assist  in  i)i'eservini.' 
its  neutrality.     Attcriipts  to  localize  the  site  of  formation  of  the  lilood 

l>iii|cins  have  not    Iieen  sui ssfiil.     There   is  some  evidence  that    (ilirin- 

ii'.'cM   is   i'ormed   for  the  nnist   part    in  the  tissues  of  the  s|  laiichnic  area 

liver'.     It  is  (|uile  possilile  that  the  hi I   forms  its  own  iiroteins.  Just 

;is  do  other  tissues,  from    the  amino  acids  it  contains. 

THE  FERMENTS  AND  ANTIFERMENTS  OF  THE  BLOOD 

'i'lic  lilood  plasma  contains  many  of  the  fcrirM-nts  present  in  the  tissues. 
Tlic  nature  of  these  ferments  has  been  the  sulijcct  of  many  invcstiiia- 
1  lulls  in  recent  years.  |)rimarily  becatisc  it  has  been  found  that  they  are 
iiiliiiiately   connected   with   the   problems  of   immunity. 

.\iiioiiir  the  ferments  the  follow  iiij;  have  been  demolislrated  in  the 
M.iod: 

!'rfi!'ii>t< s  arc   iirobably    present    normally    in    the   human    blood    serum 

ill   small    amounts,   but    they    arc    found    in    larue   amounts    in    the    white 

Muiiil   ciu'|>uscles.      A    protein    fiuciirn    to   the   bod\     if   injected    into    the 

!  ^mhI  ordinarily    produces  no  initoward   syriiptonis,   but    a   second    in.jcc- 

tii.ii   t'lillowinir  the  first   by  some  (la\s  will   prochicc  svniptonis  of  poison- 

li   known   as  (iiiiii)li  i/ldiix.     This    fact    has    led    to    the   assumption    that 

I    injcctiim   of  any   foreiy;n    protein    into   the   blood    promptly    leads   to 

'•  .ippearancc  therein  of  spccitie  pioteolv  tic  eii/yno's  which   will  diucst 

:    ■    siianirc    ))rotein    into    its   dcrivati\ cs.    which    arc    |ioisoniuis.      This 

1   '\Mr  of  the   bodx'   to   produce  specific    proteases   has   been    the   subject 

iMiicJi  research  and  debate,  and   Aberhalden  jiroposcd  a   test   for  |)rey- 

■\.  for  cancer,  and  for  other  ciuiditions  in  which  he  made  use  of  this 

i    ■  iiiiiiicnon.      lie   believes   the   presence  of   placenta    or   tniinn'   tissue    to 

'     iM'  tiic  )>rcsence  of  proteins  that  britiir  about  tlic  |)riidnctioii  of  sjiccific 

'  ■  iits  whose  duty  it   is  to  rid  the  system  of  tlu'se  substances.     <'ther 

-li^.-'.liirs  fail  !n  find  the  specificity   in  pi'ntcnlylic  acUuM  claimcil  by 

'"■  ' '■  I  lalden.  and    believe   that    proteolytic    ferments    which    arc   capable 

•'1     I'l;!  siiMi.'   fnreiirn    proteins  are   absorbed    from    the   .iliuu'iilary    canal 


J-  .M*^> 


yo 


I'Mi:  iii.iHiii    \\i>  'I'lii:  i.VMrii 


I'liiiii  ihc  (|i'.'i'sii\i'  juices  I  r.nlilviciT  I .  Sciiiic  in\  t>l  i'.'nl'irs  liiil  to  i-initiviM 
the  cliiiiii  lluil  llir  |ircp|i'iil\  lir  iMli\  ily  Ml  till' liliMid  xTuiii  is  iiiciMMsctl  umln 
tilt'  iiliiixc  coiidit  ions. 

r.lociil  ciiMliiiMs  nil  ;iMl  irii'iiiriit  Iviiiiwii  iis  iiiilitriiiislii.  'I'liis  (•.■iii  1" 
I'riiiciM'il  I'liiiii  the  liliiiiil  M'i'lllii  li,\  M'xrl'iil  MlliNtillU'cs,  iimihiim;  \\  iiirli  ;\\  • 
kiiiiliii.  cMilliiidiil    iiiiii  iiiiil  st:irc|i.     Sri'imi   tliiis  tn'iitnl  shows  slinn;.'  |>ic. 

ti'iil.vtic  iicli\it\    ami  aiiliicliycsl  ion  will  iir.      In  this  case'  there  can  l> 

MO  i|neslioli  ol'  the  s|ieeilie  oliuill  of  |>|-oteases.  Aipih'llialileii  lielieve 
thai  the  I'ellileiils  ol'  1  he  lilood  ol'  tile  pieLMiaiit  woman  me  aliie  to  diues 
the  I'laeeiital  tissue.  Iliiiiiali  plaeeiital  tissiii'  has  the  aliilit\  of  alisoil. 
illy  aiil  it  i.\  |>sin  and  it  is  \eiv  i|neslioiialile  as  to  whether  the  test  |)|m 
posed    |p\     Alideilialden    is    due    to    tile    Hew     rollliatioll    of    relinellts    or    t 

the  lenioval  ol'  the  aiil  it  r,\  |>sin  ami  the  action  of  the  protease  norniall;- 
|ircscnt    in   the   lilood. 

.\iiiliiii   I'l  nut  Ills  are  capahie  of  deeoinposiiii:  liiicleie  iiei<l  iliid   piirii 
into    the   sitiipier   Imdies. 

I.ipiisi  s  ha\e  lieeii  deliionst  rat4'<l   in   tin'  lilood. 

Aiiniliisi .     The  inesei of  starch  splitt inir  fernicnts  in   the  lilood  w.i 

llrsf  slio\>  n  i..\  .Mai;(iidie  in  1^41,  and  later  Meniard  showcil  that  y:'.' 
co'_'cn  '<]■  •■'.■.iVr'n  injected  into  a  vein  ])rodiiced  H;lyeosiiri!i.  Since  tin  : 
it    has    liceii    proM'd    conelih  i\ cly    that    diastatie    enzymes    are    iiornial! 

|iiesent  in  the  III I  and  l.vinph.     The  sonrec  of  these  eiiz.Miies  lias  irivi  ■ 

rise  to  nmcli  speculation.  Some  oliservcrs  lielieve  that  tliey  are  deri\. 
from  the  aiii.v  lopsin  of  the  pancreatic  secretion,  while  others  lielieve  tli 
they  arc  mainifactiiiecl  liy  the  livci'.  I/iLratiire  <if  the  pancreatic  dih 
is  said  to  inefeasc  the  amount  of  amylase,  while  removal  of  tlie  p;i 
creas  ma,\'   iCarlson   and   Liickhart  i    or   iiia.\    not    i  Schlesinircr )    iiieren 

the  aiii.vlasc  of  the  lil 1.      In   soiTie   forms  of  eN|>erimental   dialietes  t' 

amylase  of  the  lilood  lias  lieeii  found  iiicreas' d.  and  this  is  the  i-asc 
liiliiian  ilialietes  I  .MyciN  ami  Killiant.  If  this  is  true,  a  cause  for  l 
ilialiility  of  the  dialictic  to  store  up  i;l>  c-cc/cn  is  found.  In  impairiiir 
#if  I'eiial  function,  there  is  \isuall\  an  iiK'rcase  in  tile  lilood  amylase  ,i 
a  dcci-casc  in  the  urine  amylase.  This  has  been  su^jirested  as  liciie,' 
diau'iioslic   \alne. 

The  lilood   contains  a    feclilc  glycolytic  cn/yme  eapalile  of  dcstroyi 
ylu<-ose.     It   is  claimed  that  this  power  is  reilu' I'd  in  iliahetics  (l.epiii 

f'niiiliisi    is   I'-    iiiil  in  the  IiI.hmI  and  tissues  u.     erall>.      1(    has  the  poi' 
of  lilieraiin'.j  oxyu'en    fium   hydroycii    peio\ide    without   any   ae<-omp;i 
in'_'   o\idation    process.      Its    ]ihysiolo<_rie    siu'nilieance    is    not    known. 
is  s.-iid   th;!!    the   iimuunt    of  catalase   is   in"re:ised   'lUriuL'  cxeitemciit 
exer<'ise,    and    is    decreiised    in    conditions    where    the    liody's   acti\it\ 
lowered.      lis   detciniiiiat ion    is   cliiiicMlly    unimportant    at    )iresent. 


(■IIAI'TKK  XI 

r.i.oni):  Tin:  r.Looi)  ckkl 

r.Y  K.  »i.  I'lAK.  1.  ISA..  .M.l>. 
THE  RED  BLOOD  CORPUSCLES,  OR  ERYTHROCYTES 

'I'lii'    lllciNt    I'liiliiilii'lit    rilllctinli    of    till'    lilndd    is    to    iMIIV    iiNVircil    to    till' 

li-,Nil,>.      It    iivMs   tliis   piippcilv   rliicll\    to  tlic   icil   liloiid   cells   uliicli   ;ilf 

|iirs,iit   ill   iiiim-  iniiiiKcis  i  r).(M)(i.i'U(t  i>cr  imhiii.  oi'  l>li)inli.     Tln'sc  <"flls 

liii'  |pii-tnica\f  (iisi-s,  liiniiiir  a  (liann'tcr  of  alioiit   7."  /i.     Tlicy  arc  cmi- 

-tiiirlcii  out   of  a    fiaiiH'woik  coiiiposcil  laiiicly  of  lipoidal   material,   in 

i.     iiieslies    ol'    x\liicli    is    deposited    a    sill'stalicc    calle(l    lieiiiojfloliin,    to 

i  i.h  the  i-cinarkalpli'  osyiifii  caiiv  iny;  powci-  of  the  lilooil  is  due.     Nei- 

li.r  the  iiiaiiiicr  Ky   which  the   red  cell  carries   Its  lieiiio','loliiii   nor  the 

ihlimale   strilctiiri    of   the   cell    its<'lf   is   ac.-urately    known.      It    is   coni- 

III. .Illy   ii.-licvcd   th.at    tiic   henio^'loliin   is   hchl   enmeshed   in   a    friinivwork 

..!  stroma,  or  encased  in  the  I'dl  meiiiKranc     One  lliinu'  is  certirn'.  how- 

.  <  I  r.   that    the   union   of  hemovdoliiii    with   the  stroma   of  the   red  cell   is 

I    l;iiily  stioni;  one,  since   meie   ri-ai.'nM'ntallon   of  the  corpuscle   fails  to 

I  !ii  late  the  heuniy:lol)in.     'I'lie  fact   that  the  framewoi'k  contains  a  larire 

.Ml. .lint   of  lipoidal   sulistances  cn.iliies   the   cor|)Uscles  to   maintain   their 

-liipe   and   is   responsitde    for   tlieir  characteristic    perinealiility. 

Ileiiioirloliin   is  a   vi'i-\    complex   snlistanee   lieh.niuin^   to   the   ■j:roup  of 

''ijiiL'.'ited    in-oteins.      My   cliemieal    means   it    i-an    \>--   liroken    n|)    ;:'to   a 

pie  iilohulin  and  a   jiiu'mcnt   hematin,   containinu'  iron.      When   com- 

'   !.h    siituratcil,   oxy!.'en    is    present    in    heiiio'.:loliin    in    the    proportion 

two  atoms  of  oxvL'cn   to  one  atom   of  iron    il'i'tersi;  or  4(1]   c.c.   of 

■.  u'rii  can  he  cairicil  liy  hemotrloliin  eontaininir  one  •;ian'.  of  iron,  the 

•.•iilar  weiyht  of  tin-  m.    'cule  heiiiy:  .'iliont    Iti.tlli!!,  or  somi    multiple 

...f    I  l'.a''ci(.l't    and    I'etorsi     (see    also    p.    W'M ) .      At    this    lijruiv    the 

ill   the  molecule  would   rel)resen;    0.:'.4   Jicr  cent   of  the  total   weiirht 

ii.    molecule.     The  corpusculai'   surface   area    has   heen    estiinateil    to 

;ji"i  sipiare  metei-s.     There   is   therefore   a   very   lartre   surface  ;i\ail- 

for  the  alisorption   of  oxyirer    from   the  ;d\eolar  Jiir.   as   the   hlood 

■:  .I-].-:  jiass:   in   sinir'c   file  throuu'li   the   capillaries   of  the   Iuul's. 

■(■  tiie  jniiount  of  oxyjren  which  the  hlood  can  c;irry  de]iemls  upon 
■1.  iiio!.;loliin    content,    if    is    of    some    imjiortance    clinically    to    have 


!»2 


riii:  iiiiMiD   \M«  Tin;  i.vMni 


mii'IIhhIs  cii'  ililcniiiniiisr  iIm'  iippi'iiNiiiiiitc  iiiiiniiiit   |in'>ciil.     TIm'  aiiiniiiit 

■  if    lii'litiit;liiliiii     |ii'i'M'lll     ill    ;i     <|ll.'iMtit\     lit'    IiIiiimI     is    IInU1iII>'    tli'tniililli'il 

ciilciiiini'liii'.'illv   I'y  ciiiii|i;iriiii;  tin*  i-iilur  nf  llic  lil I  willi  stiiinliii'il  cnl 

iir>  \\  liii'li  ciiifCNiHiiid  to  kiiiiwii  n1  iciinllis  iil'  liciiin^liiliiii.  In  iinnnjil 
|nTMiiis  tin-  niiiiiiiiil  til'  liciiinirliiliiii  Miiios  iri'i'iitlx  ill  ililTi'ii'iil  iiu'cs,  ;iiii| 
in  di'ilci'  til  ili'trniiiiic  wlirtluT  nr  nut  a  triMii  lilimil  ciintiiiiis  ni.ifc  m 
less  lii'iiiiiLrliiliiii  than   niiriiia!.  it   is  iiiiprratix  i'  to  ciuisiilrr  tin-  aire.     Tli' 

'.'iralrst    \  aiiatimis  iir  lirtwri'ii   liirtli   ami   tlic  sixtfriitli   year.      At'tii 

till'  sixtcTiitli  vi'ai'  till'  liliiiiil  ill  iiialt's  usually  cniitaiiis  a  lai'urr  aiiiiiiint 
than  that  in  ri'iiialcs  i  Williaiiiscin' i.  Iiistrniiicnts  iisol  in  lii'tciininiiii; 
till'  aiiiniint  III'  ii  'Miii<:liiliiii  shiiiilil  Ih'  staiiilarili/i'i!  In  awr  thi'  valilf  in 
ijianis  liriinmjiiliin  |ii'i'  1(10  >■>■.  nl'  llniil. 

Till'  aiiiciiint  111'  lirniiiirliiliin  ^^llil•h  is  pii'scnt  in  cai'li  rnrpiisclr  in 
ti nils  111'  nnniial  is  tlii'irt'ini'  ut'  sniiii'  I'liniral  iiit'Tist.  This  rclafinn  h! 
till-  nniiilirr  uf  n'lj  nils  tn  tlir  aiiiiinnt  nl'  hi'iiniirlnliin  is  known  as  tli' 
riiloi'  iiiilix  ainl  is  cuniputrij  as  rollows:  'I'lic  a\  craiii'  iril  roiiiit  in  in:, 
is  r>,(l(Hl,tHMi  til  till  ('  iiiiii..  ainl  the  aM'i'auc  ininiiiial  amount  ol'  Ih'iiim 
U'loliiii  is  takrii  as  ]:!>'<  L'lams  in  10(1  c.r.  of  lilooij  (  SO,  Sahli;  'Mi 
Mii's.'li.T:  si;,  I'li'si-h;  ami  11(1.  Talliiuist  ini'tlinils).  Thi'sc  i-clativ 
values  ijJM'  a  riilor  iiiili'\  of  unc.  The  iii'ii-rntairc  ol'  imriiial  ri'd  ci'll- 
i|i\i<l('il  liy  till'  prii'i'iitau't'  ol'  iioniial  liciiiou'loliin  picsi'nt  jiivcs  lli 
ciiliir  ini|i\. 


The  Origin  of  the  Red  Blood  Cells 

III  I'llal  lil'r  till'  splci'ii  aiiil  tlii'  livi-r  arc  m'licrallN'  iM'lir.ci!  to  lie  ii 
spiuisilili'  iiir  till'  I'liniiatiiHi  nf  tin'  ml  lilooil  ci'lls.  In  rxtraiitcrini'  lii 
this    I'liiictiiiii    is  taki'ii   iiM'f  liy   tlii'   ri'ii   lioiic  iiiai'i'ow.      In   the  iM'iiiiiliv' 

iililiiiii    all    iril    IiIoimI    i-i'IIs   ail'   siipposcil    to   In-    liuclratcil.      In    rxli  ■ 

iitriiiii'  lilc  till'  iiiii'li'i  111'  till'   ri'.l  i-rlls  arr  lost,  ami  iioniiiii'li'atcil   I'liin 

ari'  aliiiii'  pirsi'iit   in  tlir  lil 1  slii'aiii       in   t'i'tal  lift'  am!  in  cci'tain   pail 

iilii'^ii-    I'liinliliiiiis.    till'    i;itc    of    liliiiiil    riifinatinn    is    si,    vapiil    that    smi 
iiiii'Iralril   crlls  a|i|M'ai'  ill   till'   liliiiiil,      'I'lii'   iiiiriiiiil    rcsponsi'  of  the   l""! 

til  a   Inss  of  iTil  li! 1  1-111  pusclcs  I'liiisists  in  an  ini'icascil  activity  nl'  t 

liliiiiil  rminiii'.'  cells  111'  the   icil    limii'   iiiarrnH.      It    is   not    easy    to    I'ulli 
the    euiiise    ut'   the    re'_'i'lieiat  ii  H I    nf   the    I'l'il    ciirpilselcs    or'    to    iliseii\er    ll 
iiieehanisiii  of  llnir  t'lirinaliun   in  the  lioiic  iiiannw  .  since  this  tissue  p' 
scnts   a    iiii\tiiie   111'  cells   which    are    precursors   ol'   the    \arici|   coipiise 
I'liiiml   ill  the  lijiiiiil  ami  the  iilenlily  ul'  which  can   nut    lie  iletennineil 

lieciiilly     new     iiiethoils    nl'    staiiiin;.'    Iiliiiiil     I'lir    inieroscopic    esaiiir 

llnll      have     Jiiinvveii      lllnle     llelalieil     stllll\       111     lie     Iliaiic     (III      lilC     site     .li     I 
liietliiiil    111'    liliinil    cell     riiiliiatinll        When     t'lesli    Ullflxcil    liliioil    is    tie.il 

with  siilntioiis  111'  >.iriiiiis  i|\es.  such  as  lirilliant  crcsyl   liliie,  polycliiei 


TMi:   iil.iHiii  <  I 


m 


iiirtliN  h'lic  liliic  iir  iH'iiliiil  rtil,  nil  nllifiw  isr  iiixisiiili'  ^IniiMiin'  ii|>|iriiiN 
in  sdiiif  cells  in  till-  I'lil'iii  111'  cii.iisc  uiiiiiiiliir  |iiirlirli's  i.r  lliitiiih.  wliirli 
'ji\c  a  rctiriilatril  a|>|i(':il'i'iiic<'  In  tlic  rip||iii>i'lt'><.  'riii'sr  i  rl  iiMllalfij  i-clU 
nil'  iiinri'  aliuiiijaiit  in  iiil'aniN'  lilnml  ami  in  patii'iits  Mill'i'iinir  with  sf- 
vi'i'i'  anemia  nr  lieiiioluie  jauinliee   tiiaii    in    nuiiiial    lilimil,  ami   may   lie 

laKeii  as  evident I'  llie  yontli   id'  tlie   red   eell   niiil    imt    as  a   deneiieia- 

li\e  priieess.  Siliee  llie  nnmlier  nl'  the  ret  ieillated  cells  tliat  ale  |iieselit 
ill  til"  hliiiiil  is  inure  nr  less  directly  |iri)|iiil'tiiilial  In  the  liemii|Miiet  ie 
n>'li\ities  III'  the  linne  iiiarrnw,  ennmeiatinii  el'  the  letieiilated  cells  is 
nleiinical  im|iiii'tance  in  anemias.  In  ciindilimis  in  which  animals  have 
liceii  made  |iletliiirie  li,\   the  tiansl'nsiiin  i>l   MudiI.  it   has  lieen   I'liiiml  that 

llie  nnmlier  of  retielllaled  cel|>  is  decleaNeil ;  the  liiiiie  mariuw  nl'  these 
.iiiimals    alsii    shnws    a    marked     rednctinii     in    retienhited    eiv  I  hrnlilasts. 

Till'  diminished  rate  nl'  Id I  cell  I'lirmatiiin  sumctimes  imteil  al'ler  hliiinl 

ti^iiisl'iisinns  may  he  e\|daiii('il  hy  assiiniin<;  that  the  siimiiliis  which 
.makciis   llie    I'm  inatiidi    ol'   red    cells    in    the    linne    maiinw     is   ahseiil    ur 

made  siihniirinal   nil   the   injectimi   III'   red    cells  iiitn   the   lil I.  and    thus 

till'    I'nrmaliiin   nl'   red   cells  is  dejiressed.      Small    traiislnsiipiis  are   there 

I'liie  prel'eralile  tii  lar^{e  niics  in  cases  in  which  the  rate  id'  hi 1   I'mnia- 

ihiii  is  !.M'eatly  impaired.  I>y  means  nl'  li\in<;  e  'Itiires  nl'  red  hniie  mar- 
inw  the  ililTcrent  stajics  id'  the  dcvelnpniciit  nf  the  iinrmnhlasts  inin 
tiiic  red  cnrpiisides  may  he  slndied  (Tnwer  and  llerm  i.  Snme  evideiict' 
hiis  heen  iralheied  I'mm  siieii  studies  which  pnints  tn  the  cnnclusinii  that 
ill  place  lit'  the  red  cells  liciiit:  cells  which  have  Inst  their  nucleus,  as  is 
ihc  current  teaidiinir.  they  are  rather  cells  which  devehiii  as  a  nuclear 
i'lid  and  escape  into  the  eirculalinn  as  true  red  cells.  The  nucleated 
111!  cell  and  the  red  nucleated  enrpiiscle  nt"  the  liird  are  the  pi-ndiiet  nf 
iiii  laniiclcar   activity    and    are    miirplinln^ieally    idisitical. 


Rates  of  Regeneration  of  Erythrocytes 

.Micrnsciipic  exaniinatiiin  id'  the  lilnnd  diirinu'  rapid  reiieiieratinii  nt' 
I  cells  shnws  the  pi-esenec  nl'  nucleated  fiiinis.  .Nucleated  red  cells 
'lie  liliind  have  thei'cd'iirc  liceii  taken  as  an  incvitalile  feature  nf  rapid 

I    rcseiieratinn.      The    evideiiee    upnii     whieli     tliis    helief    depends. 

ever,  is  liardl.v    eiiniplete,  since  chanues  in  the  manner  't\'  red  lilimd 

'    tnriiiatiiin    may    he    responsilile    fur    the    nucleated    rniiiis.      The    red 

■  iiiarniw  is  cniisidered  the  seat  nf  red  cell   fniinatinn,  and   it   is  true 

'   an  ahniirmal   increase   in   the  red   linnc  marrnw   usually   accnmpanies 

I   ased   red  eell    format  inn.     The  nature  nf  the  stimulus   whi.'h   lirinirs 

!i   the  lu'w   I'lirmatiiin  of  red  cells  is  imt   umlcrstnnii.     (Kyu'cii   want 

'le  an  impnrtant  factor,  since  we  lind  the  presence  nf  an  aliiinrmall.v 

'lie    nuiiiher   of    red    cells    in    colliiitions    \\here    there    is    a    scarcity    nt 


,Ji     .S: 


^1 


Ml 


Till      Ml  iNih     \\l'    TMI     I.VMI'II 


ipsv_'"'Ii   ill  lln'  iMs|iiitil  nil'.  ii>  ill   lit'f  lit   lii'.'li  iillitinlis,  nr  n  ilil1liMilt\    ii 
iti  iili>iii|itiiin   iliruiii'li   till'   liiiiirs.  iis  in   rniiL'i'nitnl   lii'iit   iIIsciim'. 
Till'   It'll   i-rlU   prniliii'iil    ri>llii\'.  iiiir   Ih'Iihii':  iiiiLTi'   ami    in   siiii|ili'   iiiiriiii 

I'lilililill  li'ss  tllilll  till'  lliil'llllll  ,'lllliillllt  III'  lli'liinuliillill,  lillt  tlirit'  Nllilpi'  illi' 
si/I'  nil'  lili|)rii\ilii,lli'lv  liipllllill.  mill  Irw  lllli'lriitril  rrlU  ill'i'  |>rrM'lll.  I 
till'    ii"_'i'iii'iiitiiiii   111'  fi'il    I'i'lls   wliii'li    i>    riiiiiiil    ill    pi'i  iiii'imis   .iiii'iiiiii,    \ 

Mini    til lU    ■■iilit.'iiiiiiit;    iill     lllillMliill V     liil'U'i'    iiliiiiillit     III'    lirli|i>i.'liilMi 

'I'll;'    III!   I'i'lls    ill    tills   ilisi'iisi'  Ir         iilitiiii'iiiiil    rmiMs,    iiijiiiv    lii'ini;    1,'ir'ji 
with    III'    witliiiiit    .'I    iiiH'Iriis,    iiii.      ciiiitiiiiiiiii;    liiisii-    slniiijnu'    ui'iiinil 
This  t\] I'  Iilnml  imII   I'liriniitiiiii   is  ilin-  In  i|i'i;i'iii'i'iiti\t'  rli.'iii'.'i's. 


The  Fate  of  the  Erythrocyte! 

Till'   Irllirlll   111'   lilV   111'  till'   I'l'ij    111 I   ri'.i    is   l.liiuiiiw  II.       I'',s|i|ii;iti's   li:i,' 

lljtcill  till'  iliiil)  cNi'liliiili  111'  liilr  p'lrnii'iits  ;il'i'  lint  li'linlili',  siiiri'  l|iii>|. 
mill  \Vlli|l|i|<  llJIM'  sIlllUII  tlljll  till'  |li!.;llll'llls.  ill  |i:|l'l  ill  Irilst,  .'ll'isf  I'll. 
|ii'.'llli'llts  wliirli  till'  ii\i'r  lijis  Miiiili'  ill  I'M'i'ss  III'  its  lli'i'ils  fill'  till'  111,11  ■ 
I'.'li'llll'r  111'  licli|iii.'iiiliili.  iiinl  wllirii,  lint  lirilin  llcrili'il,  iili'  rMM'rli'ii 
Tlii'ii'    i-     'II    i|ii>stiiiii    ImwrMi'    tliiit    I'M'i'v    ci'.x  tliini-ylt'    siiiuiiT    nr    l.i'' 

ilinli'i'i.' lisiiitiui'iiliiiii.  ;i    |iiiii'i'ss    liiriiinly    tliiiiiy:lit    to   lir   iisIhti'iI 

li\    till'  iinri'slimi  of  till'  ri'il  III I  rill  liy  a  pluiu'i'i'vti'  in   tlir  s|ilri'ii   > 

in  !i  lii'iiKilviiiiili  L'lmnl.  till'  liiiiiii'.''|iiliiii  nf  lln>  ilisinti'irr.'iti'il  im'II  lii-iny  - 
liii'  mill  ciii'iiiil  til  till'  li\"",  "1  11'  it  is  )  inki'ii  lip  iiiti)  lifniiitiii,  wlii' 
till'  liiiily  stnri's  I'nr  I'litiiii'  iisi',  aliij  into  liilr  piniiii'iits,  wliii-li  mi'  ■ 
rrctril.  Ii'iills  iiml  Ui.lii'it-iiii'  I'.iil  li>  liinl  cviilriiiM'  llijif  tliis  )iiiii'' 
iM'iMiis  ill  Minn  til  nil  i'\1i'iit  siitlirii'iit  tn  ni'i'mint   I'm'  tin'  iMniiinl  lirstMi 

tinii  III'  till'  111 I  rilU.     lliiwcMM'  lliry  linvi'  I'l'i-i'iitly   I'miiiil  nimtlii  ' 

iiiisnspi'i'ti'il     nictliiiil     I'nr    Iili.ml     ili'stnictiiin     in     nil   miiiiinls   thus 

sfinlird      iimiu'ly.  tlii'  ilisiiiti'i.'r.'itiiill   nl'  tin'   lil I   rrlls  liy    t'lnirMii'litnl  i 

\\liili'    llii'\     nil'    i-iii'iil;iliiiir.    withoii!     hiss    ul'    thi'ir    li('iiiiii;li.liiii,      Tl' 
I'lnirniciiti'il  nils  nil'   tniiliil   limst    I'li'iini  lit  ly  in  tlir  spli'cii.      They   lii'li. 
thnt    till'   suinll    ill  I'liinii'il    I'i'lls,    kimwii    ns   niii-i'iiry  ti's   ninl    pnikilnrv  • 
iilisci'x  I'll  in  si'M'i'i'  I'Npi'iiiiii'iitnl   mii'iiiins,  ,iri'  ilin-  imt   tn   tlii'   I'.irt   t- 
tlii-y  ;iri'  piniliii-i'il  liy  the  liniii'  inniinu.  Imt    rathi'i'  In  llic  ('net   thai    ' 
Mimi'nw  ill   iis  niiriiiii'  rnmlitinii   is  lint   nlili'  tn  proiliii'o  n   I'i'sistniit   • 
tlii'iicyti'    ami   rrairiiiititatinn  tlii'i'i 'fnri'  takes  jilarc  tun   nailily.     A    ■ 
ilnr  ciiiiilitinii  may  I'xist   in  tin'  m'mtc  aiii'inias  of  man  ami  arcniiiii 
till'   U'l'lli'ial  lliull   I'csistmirc  nl'  the   rril  cells   I'lillliii   ill   the   liloml   nl'  1 1 
pnticiits.  innsnineli  as  the  wenk  eells  are  ireneially  I'lau'ineiiled  vei'\   ^ 
nl'ti'i'   they   ni'e    fniineil       l.iiinr   aL'n    Kliilich    stateil    that    the    niieini- 
mill    J)iiikilneytes  nf  aiieillia    me   the    lesiilt    nl'   t'laKlllelltatinn    nl"  the   i 
in    the  eiii-lilatiliu'  Iilnml,    Imt    lie   lieliev  eil    lliat    this    frairilieiltalinll    w.i 


'I  III     MI.OIIIl   II  II, 


111  pllll 


till     <l 


niMiiii    III   (III 


l,r   I 


II    inrlriisc 


till-     tnllll     sllirilCl'     III'     till'     ri'il 


.1  llx.  Tin  lllfiniiltr  filtr  111'  tlir  li'il  itII  t'liiL'llKlits  in  iml  Ixlmwll.  It  is 
.iMsiiiiiilili-  III  silli|>iisc  lliiit  til"'  riMuiiH'iili'il  liits  riintiiiirna  h<iiii>ulnliiti 
III'    ('Mirii'ii     to     till'     llMT,     ^lirrc     the     lu'lllDulnliiM     is     t  I'llllHl'nriili'il     illtn 

lii'iiiiitin  :iiiil  liili'  |ii<.'iiiriitM. 

Hemolyiii 

Alintili'l'    lllitlliiil    III'    nil    liliiiiil    c'i'll    ili-'ll  ili'liiill.    wliii'll.    linwrMT,     Imi 

iioi  liikf  |)liii-('  iiiii  iiiiillv,  in  liy  lii'iiiiil>>is.  Till'  iiiituii'  III'  till-  ciiiiiliiiia- 
linii  III'  till-  lii'iiiiii.'liiliiii  with  till'  stiiiiiiii  iif  till'  li'il  nil,  iis  iili'i'iul.N  n- 
111,11  Kril,  !•<  Mill  ili'liiiitrly  kiiiiwi  .  'riiiil  it  is  mil  iinTi'ly  I'lititiiiliril  in  a 
^,ic  is  nIhiwii  liy  till'  I'iii-t  tliiit  tlic  i-rll  iiiiiy  In-  cut  iiitii  Mis  witlimit  tlii' 
111  iiiiii;|iiliiil  liiilivr  sit  I'ltr.  Ill  siiiiii'  inaiilii'i'  tlli'  liiliiii'.'lnliiii  is  i-lirl'l- 
ii-;illy  liiiiiinl  \\itli  till'  stiiiiiia  nl'  tin'  ml  I'rII.  I'lmii  wliiili  it  I'aii  l'<' 
111  111  liK  a  liliiiilx'i'  III'  |ili\  siriii'liriiiiiMl  ainl  fliiiiiii-al  airi'iits.  'I'liis  |iiiii'- 
1 -N  is  kimwii  as  III  iiiiil'isis,  iiiiil  till'  miIisImiu'i's  wliirli  l.iiiiu'  it  aliiiilt  an' 
kniiwii  as  liiiiinlv  I  ii-  airciits.  Till'  niaiiiHi'  in  wliirli  tlii'si-  aui'iils  rlVi-i't 
ill'  I'l'li'asc  III'  li('iiiii;.'li>liin    I'liilii   tin'   lilnml    is   qiiili'    vaiii'il. 

If   till'   lisllllltir    IH'Issllfr    III'  till'    |llas||l.'l    is    liUMTliI    li\     ilihllinll.    till'    |iirs- 

■MIC   uilliin   til ir|iiisi'li'   ri'iiiaiiis   liiuli,  ainl    watiT   is  alisniln'il   li,\    tin- 

'■1  'I     If  tliis  alisi>i'|itiiiii  is  Miriii'ii'iit.  till'  t'i'll  iii|itiiri's  :iiii|  till'  lii'iiiiii:liiliiii 

i~  ilisrharu'i'il.     l''iir  lliis  if::;-iiii   it    is  ii ssai'v   in   ililiitiiiir  tlic   li! 1   In 

,,.•   snliiliniis   111'   suit    li,'i\  ill'/   an    iiMiiiilir    |iirssii?i'   ciiual    In    tli;it    nf    tin' 

1  III  pi'iiti'i't  till'  iTil  I'i'll  rtniii  lii'iiiiil.v  sis.     Tliis  is  iilitiiiiHil  liy  iisiiii; 

'lit    |ii'r   rent    siiliilii.il    .it'    siiiliiini    clilmiili'.      i'nlti'i'    ii'siilts    air    liail, 

'''■'.  i'\  IT.   1>\    usiiiiT  I'itliiT    i>iiii_'i'r's   siiliitiiiii    ili.M    |iiT   I'i'iit    Nai'l.   (i.irjii 

i!  iTiil  t'at'l .,  ami  O.li:!  piT  niil  l\<'b  m-  l.iii'ki''s  snluiiun    nil  piT  i-ciit 

^  '1.  (in-JI   |it'i lit    i'arl,.  iiiiVJ    piT  I'i'iii    Kt'l,   11.11)  ii.ii:!    \h'\-  ci'iii 

Mill    ami  111    JM'!'  ('ciit    jtliirdse.i. 

'  >    iiniiiial    I'liipiisi'lis   lii'iiinlysis    iii'ciiis   tn    a    sniiill    I'vtiiit     in     snlii- 

I'liiitainiiii:   alimit    ll.4"i   piT   ci'iit    nl'   sinliuiii    ililin'iili'.      In    t'citaiii 

>  till'  I'latrility  nf  tlir  rinpiisi'li's  mav   In'  iiii'ii'asi'il  ijintli'i'*. 

i  i      iiH'iiiliraiio   ami    struma    nf   tin'   iTytlii'iicyli'   ciiiilaiii    lipniilal    iiia- 

!'  .i!    \\liii-li    is   siiliilili'    ill    alriiliiil,    ctlii'i'.    fatty    ai'iils.    ami    liili'    salts. 

\i'liiinii  III'  tlii'si'  ai;i'ii1s  to  till'  liliiiiil  liriiiirs  alimil    lii'iiinl.v  sis,   prcsmii- 

''    liy  ilissiilviii!.'  till'  lipniilal   liiati'i'ial   pri'si'iit       Tin'   Iniiinlxsis  wliirli 

i's    uiili   sapnnin    is   similar    in    typi'.    siiii'f    sapmiiiis   (•niiiliinc    with 

'  'lis   till'  riiiii|)iiiiml  licinir  snliililc  in  \v;iti'i'. 

Ill"    lirmnlytic    i>rnp('rti('s   of   sitiiiii.    wlictli    ■    -liry    .'iio    I'miml    tn    In' 

ii.ill\    pri'siMit   wlitn   till'  lilnoils  nf  rci'taiii       ,imals   aro   mixi'il   <>v  In 

i'i'"'lii 1  ai'tificiiilly  Iiy  llii'  in.ii'ctinii  of  fi.i.iu-ii   ii'il  crlls,   fiiiiiisli   ,'i 

;  i.jri'l   nf  irri'.'it   iiiti'l'i'st    fi'iiin  tlio  stainlpoint    Imtli   nf  iiiiiiiiiiiiiliiL:y   anil 


!m; 


■nil:   iii.iHiii  A\i>  Tin;  i.vmi'I 


111'  rliiiir.il   iiicdii'iiic.     Tlic   1icmiip!\  t ii-  scnmi    prtiiluci'd    liy    the    in.jt'cliui 

111'  riiti'iu'll  ciillillsrlcs  owes  lis  .MfllNilV  I"  two  NlllislillUM'S.  Tilt'  iilh 
imIIiiI     lllc    iliiihiiii  jihir,    111'     illlMllllli'     IkmI.V.     is    spi'cilic     il^ililisl     till"     t>  |i 

111'    cell    in.ji'cti'il    mill    is    iiiiTi'.isi'il    iliuinn    iiiiiiiiiiii/iitinii.      Tlii'    sccim. 

Iiiiily    is   till iii|ili'iiii'iit  ;    it    is    iHiiis|(t'cilic.    nini    is    not    iiicri'.iscd    ilm 

iim  iiiiiiiiiiii/.'it  imi.  ( 'iiiii|il<'iiii'iit  is  ili'stniyi'il  liv  licatiiii;  tlic  scriiiii  !"' 
uiii'  limir  lit  .').">  ( '.,  Ii',i\iim  llic  iiinlmci'ptur  mIiuic  iiri'sciit.  ( 'orpiisclr^ 
plin'i'il  ill  siii'li  si'i'lliii  iiic  imt  lii'iiiiilv/cil  until  (■iiiii|ili"ii  'lit  eitlii'l'  IVhh! 
IVinIi    iiMiir.iiii'  III'   I'i'iiiii   niiiiiiiiiiiiiiii'  si'iniii    is  ikIiIciI. 

Till'  srniiii  111'  ;illilii;ils  |iiissrssili>i  llJit  IIIJll  llcliliilv  t  ir  prii|>i'll  ics  tuVMIIil 
tilt'  i-iii  piisi-li's  111'  iitlicl'  ;iliilii:ils  liki'wisi'  uwcs  its  rtTri't  tii  the  jniiit  Jii'tii.i 
111'  .'Itllliiirrptiirs   ailil  riiliipli'lili'llt. 

( >riliii:iii!>  tlic  si'i'iiiii  I'luiii  iiniiinils  uf  mic  species  dues  lint  cxITiIm' 
liciiii>l\  tic  pi'iipei'ties  til  liloiiil  I'niiii  iiniitlicr  iiniiiiiil  ul'  tlic  sniiic  spccii  v 
III  iiiiiisiimI  ejises,  liiiwcxcr.  tlic  scrniii  ul'  iin  iiniiiial  will  iirmliicc  liciiini 
vsis  111'  111!'  ciirpii-i'lcs  111'  an  aniiiial  nt'  llic  same  species.  Siicli  sera  a" 
said  to  |iiissess  isiilieiiiolx  sins.  The  fact  is  nt'  LTieat  iliipoitance  in  tj. 
ti'anst'iisiiiii   oj'  liliiiiil   I't'oni   niic   iiidi\  idiial    to  aimtlier. 

Till'  cause  of  the  acute  hcinolysis  which  occiiis  in  tin'  disease  paii'\ 
Nsnial  hciiioLiliiliiiiili  ia  is  not  known.  It  is  |)iolialily  due  to  the  picscm  ■ 
III'  :i  hciiinlvtic  siilistaiice  which  unites  with  the  Mood  coipnsclcs  ;;' 
teiiipeiatiii'cs  lieiiiu  the  iioiiiial  limly  teiiipcratiire,  silK'c  the  attack  h- 
lows  e\piJMiic  to  cold,  and  lilnoil  I'loill  )iaticiits  sllli.ject  to  the  coliilili'.  i 
iiia,\    lie  hciiioly /I'll   /'//   rilm  liy  ciiolinjr  and  siilisei|iii'ntly  heatini^  it. 


LEUCOCYTES 

There  are  a  nniiilicr  uf  \ai'ieties  of  white  cells  in  the  lilooij.  These  ; 
ilitTci'ciitiatcd  from  one  another  liy  their  shape,  slaiiiilii;  properties.  :\< 
the  irianiiles  in  their  prutoplasni  We  nia.\  divide  them  into  two  in;: 
L'loiips     iiniii;raiiiilar  iiHiiioinich'ar  cells  and   '.'I'aiiiilar  iiolyniiclcar  >■•■ 

The  iiiiii'jraiiiilar  iiionnnnelear  cells  arc  tcriiied  hiiiiiilnifiili  s.     'Ywn 
rieties  arc  ililTcrciilialcd,   the  small   and   the   larce. 

The    ■-lliall    liioliiillllclcar    leucocyte    makes    up    Iroiii    "J.!    to    'JS    per    «■ 
iif  the  total  lenciic.\  lis  and  the  lai'LTe  mononuclear,  from  '2  to  4  per  < 

'i'iie    piil\  nuclear   leiicncyics   are   dixiildi    into   three   "jnuips   acioi'' 
to  ■.\lictlicr  their  uraniiles  slain   with   basic,  neutral   or  acid  stains.     '! 
Iciicucyles   that    slain    with    hasic   dyi       i.r   the   hasophile   cells,   are   \ 
few,  niaki'i'j  up  less  than  one  per  ce        if  the  total  count.     J.iUcwi^' 
acid  stainiiiu    maiiiilar  cells,  acidop'      •,   are    few,   coniprisiii'.;    from   - 
4  pel    cciil  of  the  total  count.     The  i     ist    r  rous  are  the  iiciitiopk 


•nir.  iii.oon  <i;m. 


97 


,,r  the   piilynuclciir    Iciifocv  Ics.    with    iiiMitriil-stjiiiiiii'r    ■.'liiinilcs.      Tlit'sc 
iniiiprisi'  t'roiii  <).">  to  7r>  pci-  cfiil  ul'  tlif  liiliil  I'lunit. 

Aiiotlicr  tvi)c  tif  wliili'  cell  is  kiuiwii  iis  tin-  t riiiisitioiiiil  cfll,  hcciuisp 
it  was  s'i!.,i  "(1  to  represent  an  inlernietliate  t'onii  lietweeii  tlie  iiicuiii- 
,111,1  )  '.iiin ';■;'!■  ,.  !!s.  rioliably  sueli  t fansitiiHis  dn  not  neenr,  and  tlie 
trans  !inn:il   Iniei  i-.\  t  ■    s  related  to  tli'   iiionoiniclear  cells. 

Th      |i  :,^  nii.le.n-       ils   oriuinate    in    the   Iwine    niarniw,   an<l    fur   this 

reason    ii,;vi-   1 n    t.  rnied   myeloid   cells.      Tiiey    develop    floni   cells   in 

the  hone  niarrovv  .eiiiied  niyclohlasts,  which  are  noiijrranulai'  and  con- 
tain a  larfie  inndens.     In   tli nise  <>(  dcMlo|)nu'nt    the  characteristic 

'jrannles  apjicar,  aiMl  the  nncleus  remains  ronnd  ami  later  heconu's 
l.ihidatcd.  These  intermediate  forms  are  called  myclo<-yt.'s.  Tlie  nnmo- 
hu.'lcar  cells  ori-rinatc  in   the  lymphatic   tissues  of  the   liody. 

The  leucocytes  i)osscss  the   ahility   to   make   ameboid   niovcnwnt    and 

to  infrest    forciiin   parti<des  \\liii'h   may   he   presented   to  them.     On   ac- 

.ount  of  this  latter  ahility   they   are  commonly   called  jiha^'oeytes.      In 

iIm'  process  of  inflammation   the   liMleocytes  assemhle  at   the   spot   uhich 

I-  till'  seat  i>(  the  iiijnry  or  infection,  and  reiiM)ve  the  foreiirn  or<;anisni 

!■  necrotic  tissue  hy  insiestini;  ami  ditrestinir  it. 

It    is   not    definitely    known    whether   or    not    the    lymphocytes    fmn-- 

tiiii  as  i)ha^t<icytes.     Othei-   functions  hesides  those  as  jihaijocytcs  have 

i..  m    aserihed    to    the    white    cells,    lint    tlii'y    are    imt    universally    ae- 

■  I'tid.      The    iiumlier    of    leucocytes    in    the    lilood    is    snli.jcct    to    con- 

iriahlc   variation.     They    normally    numlier   lietween   (i,(HI()   and    S.ltltO 

I-   e.nim.      At    the   liei-ilit    of   dijjcstion    and    after    strenuous    exercise 

rr    is    usually    a    small    increase,    and    umler    iiatliolo^dc    conditions. 

■iallv    in    infectious    diseases,    this    lieeomes    (luite    marked.      Some 

iuiis    inci'case    the    |)olym(irphoiinelcar    cells,    while    others    add    to 

phoc.vtes.     The   factors   •rovcrnini:  the   t.\pe   of   increase   arc   not 

■un,  nor  are  the   functions  of  the  various   fmiiis  differentiated. 

The  Blood  Platelets 

TImm'  are  small  oval  particli's  alioiit  -^  n  in  diameter,  which  are  fnnnd 
ii!  MiL'i  nnmliers  ( ■jriO.ddO  to  the  c.mm.i  in  the  lihiod.  They  arc  sn|>- 
p.ivrd  to  lie  formed  from  iiarticles  of  iirotoplasm  which  are  pinched 
"■V  from  ihe  larjro  Mood  cells  in  the  lione  marrow.  Their  hioloiiical 
iiid  rheiiiical  properties  are  not  understood.  They  proluilily  iday  a 
''!>    iiiipoitant    role    in    tli laiiulation    of    the    Idood    (see    paire    l<i:>). 


',Jgj^ 


CIIAITKW  XII 
I'.Lddl):    I'.LmoI)   ('l,()TTI.\(i 

On  li"ii\ilii:  llic  lilood  vessels,  the  IiIoimI  clots  Sd  Jis  to  t'ol'iii  a  pluL', 
wiiicii  assists  in  |ii'e\  eiitini;-  tnrtlier  lieniorilia're.  The  elollini;  inii^l 
theicl'ofe  lie  ccinsiiliTeij  as  a  piiiteet i\ c  in<'i'hanisni  ajiai'"^'  exeessi\i 
<ii-ainiii^'  ot'  hlooil  ont  of  the  oisranisni.  When  the  wouiiijed  vesseK 
ave  small,  the  elottintr.  alonir  with  const  lielion  of  the  daniaired  vessel-- 
and  the  foinialion  in  theiii  ol'  throinlii  containiii>;  lai'<;e  iiiiinliers  oi 
l>latelets,    serves    to    elTeet    complete    stoppai;e    of    the    lieinoffhasri'    excii 

tlioiiu'h    the    hi I    piesMii-e    may    not    have    hecome    mateiially    redneed 

The  irieater  loss  of  lilooij  from  larirei'  vessels  causes  the  aiterial  ])ressni'e 
to   fall,  and   this  enahles  the  clot    to  stiiVcn   and   seal    the  Avouiid   hefori' 
the    pressure    aiiain    rises.      When    the    clottiiifj;    ])ower    of    the    lilood    i- 
snlininiiial,    lifi'    is    eiii!any:ered    h\'    even    trivial     Wounds;    ninier    the^i 
conditions    the    siiiallest    surface   sciatcii    may    coiitinne    to    lileed    e\ce^ 
sively   in   spile   of  whatever'   local   treatment   is  a|i]>li<'<l.     The   nuist    e\ 
Ireiiie   deiri        of   this   condition    occurs    in    hemophilia,   a    disejise   whidi 
is    cliaiacteri/c(l    liy    a    most     interestini;    family    hi;  tory     namely,    th.il 
althoiiLrh    it    alTeiMs    only    eertjiin    of    the    male    memhers    of    a    famit.^ 
vet   it    is  transmitted    from   •rcMcration   to  ixcneration   liy  the    female  sid' 
alone.      The    disease    has    e\i>led    in    certain    id'    the    royal     families    '■' 
Kurope    for    many    L'eUcr.itions,    vvhiidi    has    maile    it    ])ossilile    liy    cm 
snltinir    the    L'erie;ilouic    trees    to    demonslrale    the    infallihilily    of    tli  ■ 
law    id'  inheritance. 

The  clottinir  of  the   hlood    is  also  either  depressed   or   increased   in     ; 
\;iriet.v    of   plivsioloyic   and    i>atholo<ric   conditions.      We   siiall.    hovvcvi    . 
del'er    further   consideration    of   these    until    we   have   learned    soiiicthii 
of  the  naturi'  of  the  factors  which  aie  res|)onsihle  for  the  process  ilsi' 

The  Visible  Changes  in  the  Blood  During  Clotting 

In  a    few    nnnutes  aftci-   it    leaves  the   Idood   vessels,  the  Idofxl   t'onii- 
.jelly  like  clot,  v\hich  adheres  to  the  walls  of  the  container  in  which  i 

III 1    is    cojlcited    and    soon    hecomes    so    solid    that    the    vessel    may    i 

inverted  without  spilliriLr  iii\\  of  tire  lilood.  ('lotting'  is  now  said  to 
complete.  The  'lot  soorr  lieiiins  to  eonli.'icl,  and  as  it  does  so,  drops 
clear  Itnid   or-  serum   liecorne  expressed   and    lloat    on   the  snii'aee  of  t 

!is 


Iii.oni"  <  i.o'niNd 


00 


flnt  or  (•(illcct  ln'twccii  it  iiiiil  till'  Willis  lit'  the  ciintiiiMci',  so  tli;it  lifter 
soiiii'  time  till'  cliit  Iii-i'mUs  MUMy  I'loiii  i  lie  I'oiiliiilH'f  Jiml  cuiiics  tci  tlnjit 
ill  the  sciimi.     'I'll''  JjittiT  iiiii\    111'  pi'ijVi-tly  cIcMf.  Imt   iisiuillv  is  iimrc  oi' 

Irss   ii|i;il('sci'iit.    piiiH.v    1 iiusc    lit'   tiic    picM'iii'c    111'    fat,    iiiid    jiai'tly    lic- 

railse  111'  li'ili >l<'-^  w  liirli   liavr  inieiati'il   nut    nl'  till'  I'liit   111!   ai'ciiiiiit    ui' 

tlirif  piiut'l'  of  liiapi'ili'sis. 

If  a  (iinp  iif  ffi'slily  slii'il  liliiiiil  is  I'xaiiiiiii'il  iiihIit  the  iiiiiTnsi'iipi',  it 
will  he  (ihservci!  that  the  first  step  in  eliittimr  i-misists  in  the  f-.i  niatinn 
of  line  threails  railiatiiii:  fnnii  fuel,  whieli  are  uinhiuhleilly  the  lilmiil 
platelets.  The  line  threails  are  ealleil  lilirhi.  They  multiply  rapiilly, 
su  as  111  fiiriii  an  interlaeinii  iiieshxMirU  v\hieh  eiitaiiirles  the  reil  liliiml 
i-iirpiisi-les  ami  lelleiiev  les.  I'.v  the  Use  <•(  tile  lilt  ralllieli  iseiipe  I  pa!.'!'  .'i^  ) . 
Unwell'  anil  nthers  have  nhserveil  tha,  the  lihriii  (  prmliieeil  hy  aihliii'-' 
thrniiiliin    In   ii\alatei|    plasma  i    is    reallv    ilepnsileil    in    the    I'nriii    i>\'   line 

erystalline    iieeilles      "lilirin     li lies"      uhieh     lierniiie    paekeil     tnyelher 

as  the\  inerease  rapiillx  in  nuiiiliers.  Althnuijh  the  prneess  nf  elnttinu' 
eniisists  therefnie  in  the  enuversinii  nf  a  li.\  I'.rnsnl  iiitii  a  hxilrnyel  i  see 
pajre    (Id),    it     is    a     Ulliiple    prneess;    a     snlutimi    ti\'    tin-    lilnml     prnlein    \\liieh 

is  respniisilile  fur  the  fiiriiiatinii  nf  the  lilirin  i  lilniiiiejen  i  may.  liNe  ntlier 
enllnjilal  siiliitinns,  I"-  preeipitaleil  in  a  \ariet>  nf  ways,  hut  it  is  mily 
when  the  eniiilitiniis    ire   favnralile   fur  lilnml  elnttimj   tha'    tilniii    neeilles. 

ami    therefore    lilirin    threails.    are    d-i^ 1.      The        -inil    ,<(    iin  ertehrates 

rnriiis   a   stnietlireless    eel    wiieli    it    elnts    I  Unwell  I. 


Methods  of  Retarding  Clotting  of  Drawn  Blood 

Til  iimlerstaiiil  the  nature  of  the  elntliny  pn.eess  ami  the  faetnl's  that 
are  respoiisihle  for  its  neeurreliee,  it  is  ail\  .ntaueniis  tn  simplit'y  ihc 
ennilitimis   sniiiewhat    liy    eettini.'    riil    of    the    leil    empuseles   ami    must    111 

the    ntlier    fniiiieil    elements    of    the    hi I    anil    lluli    llsiiiLr    the    lliiiil    in 

whiell    these    are    sUspeliileil    in    liviuu    lil I      liaiiielx,    llii     iihlsiiiit.      This 

reparation  of  lilimil  into  eorpiiseles  ami  plasma  is  reailily  etl'eeteij  either 
liy  si'diiiieiiiation  oi'  liy  eenlrifiiuine  after  measures  have  heen  taken  1i> 
inliihil  or  lil-eatly  ilela.v  the  elnttin!.'  proeess.  The  melhoils  liseil  for  this 
purpose  are  iiumernus.  A  few  nf  the  must  important  are  as  fnllnws: 
'I'  Keeping:  the  hhinil  at  a  temperalure  \  er>  sliu'htlv  almve  free/iliu' 
point.  This  melhnil  is.  hnwever,  not  very  el1'eeti\e  unless  the  lilnml  is 
iiiiiiieiliately  feeei\i'il  intn  narrnw  vessels  plaeei!  in  iee  ami  the  lempera- 
'ure  kept  niiisl  strietly  at  the  low  level.  In  the  ease  nf  lim-ses'  lilnnd  anil 
ntlier  slow  ly  clottine  hlooils,  the  methoil  siieeee.ls  without  these  |treeail- 
tinlis.  Cji  Ueeeivine  the  hloiiil  <hrnuuh  a  strietly  eleail  ailij  smnnth  eall- 
liula.  ei.aleii  with  a  la.ver  of  paral'lin  m  \a>elilii-,  ililn  a  Xi-sel  .^imilarlv 
'■oateil.     This  metlinil  is  of  praetieal    imiiorlanee  when   it    is  iieeessarv    to 


d 


.Jf-.t/"'' 


it 


I 


1011 


■nil.    lilnuD    \\i)   Tin     '  VMI'II 


ti'iiiist'iiM'  Mi'i'il  williiiiil  inakiiii;  ji  \cs>  \  <'ss('l  iiii.-islciiiiosis.     (i!)    .Mix 

iliu:  till'  IiIimkI  \\\]]\  clifiiiicMU  lliMt  lire  ..ipnlilr  of  I'dimv  im;  tlu'  cillciillii 
iVdiii  sdlul  mil.  Siii'li  iciiirciits  ;iit'  |»iiliissiiiiii  or  sniliiiin  uMihitr  I  in  :i  cuii 
i'iiitr;itiiiii  of  (1.1  i)i'r  cent  ;it'ti'r  niixiiiui.  niiil  sucrniiii  ttiicnidi'  iiml  sodiuin 

cillMlr  I  ■_'  prl-  cc'iil  sdluliiill,  \\itll  olh'  |);ill  dl'  till'  Millltiull  tii  I'lHir  liilll-- 
iif  liliHHJi.  (4'  .Mi\inLr  till'  IpIimhI  with  iTitiiiii  lu'iil  i;il  siills,  piirticuhiil.v 
tile  .sill|iii,iti's  111' sdcliiiiii  iiikI  iiiiii^lltsiiiiii  (one  |i:ii't  iil' "JT  ])('!•  cent  siiliitii'ii 
dt'  liiii^iie^iuiii  siil|iliMle  liiixeil  willi  t'lilir  |iiiils  nf  lilciodi.  Uliiinl  tliu'- 
tieiileil  is  kiKiwii  as  "salted  lilciitd."  and  tlie  plasma  separated  liy  ceiilii 
I'liiriim,  as  •'sailed  plasma."  t  'lot tiny;  is  reaililv  induced  liy  adding;  w  at'  ; 
to  llie  sailed  Mood  of  pla--iiia,  and  in  this  \\  a \  diiiiinisliin^  the  concei; 
tiation  of  till'  salts,  i .")  i  The  addi'ion  to  lilood  ot'  one  of  a  class  id'  slih 
stances  known  as  antitlii-oml>iiis.  I.eecdi  extract  or  the  iiniilied  snlpstaiie. 
separated  from  it.  known  iindi'r  the  trade  naiin'  of  "hirudin."  and  siil> 
stances  prcsi'iit  in  Mood  removed  from  animals  after  they  have  lieei, 
injected   with  jieptoiie  solutions,  are  examples. 

'I'he  metho(U  which  have  Just   lieell  desciilied  are  those  api)lied  to  Moo  ! 
after  it    has   left    the  lilood    vessels,      .\notiier   interest  ill;;   u'roii|)   of  ami 
coau'illants   pre\eiit    clottim.'  mihi   irlnii    liijii-li<l   inlii   Ihi    hhiiiil   rtssils   ..; 
///(     liriiK/    iiiiliitiil.      '{"he    most    |)o\\erfnl    e\aliiple    of   this    uroll])    is   sliak' 
xeiioni,  I'crtain  \arieties  of  which  can  |irevi'iil   clottiii'.r  in  the  dosa<.te  i  ' 
',,,11  "f  a  iiiillii.'raiii  for  each  kilojiraiii  of  liody  wcinht.     Siiiii|;ir  Imt  niiicli 
less   iioteiil    effects   ;ire   |)rodiiced    liy    the   in.jeclion   of  several    ]>rotcolyti  ■ 
enzymes.   Imt    most   attention   has  lieeii    jiaid   to  the  elVect    <d'  coniinercia' 
liejitoiic  injected  ill  solution  intravenously  in  the  proportion  id' O.o  ^rani 
to  each  kiloj.'ram  id'  liody  wi'ii.dit.     Ulood  snlisei|nently  removed  ii|)  to  alioi, 
iialf  an  hour  or  more  does  not  clot,  jiinl  as  we  have  alrcjidy  seen,  if  ;iddi 
to  liloorj  from  another  animal,  materially  retards  clottinir.     This  ^'ronp  ' 
//(/;•(/  rHiiiii  aiitico.i'jnlants  is  iiarticiilarly   iiitere>tin<.'.  since  none  of  il 
sulistances    lieloiiirin!.'    to    it    is   e;  palile   of    prevenlin^'    clotting'    id'    hlo. 
v\  hen   mixed   w.ith   this  after   it    has   lieen   shed.     Tlicii'   ai'tion   thercl'" 
olivioiisly    depeiols    on    the    ]iroduction    of   some    snlistance    in    tlie   ho.l 
prohalily,  as  we  shall  see  later,  in  the  liver,  since  th(\v  f;iil   to  act  ,'ift' 
the  removal  of  this  or'_Mn  from  the  circulation   (sei'  piiire  ]]\). 

Tli<  liiiii  of  chilliiifi  varies  u'reatly  accordiiiu'  to  the  conditions  iiin! 
whicli  the  lilood  is  collected  ami  the  animal  from  \vliich  it  is  derivi 
Human  Mood,  for  exiimple.  received  into  a  test  tiilio  from  a  piincli. 
thi'oilu'h  the  skin  nia>  clot  ;it  aiiv  time  within  three  or  ten  iiiiinites.  V 
minutes  Iieinu  rakeii  as  ;in  averaL'i'  time  for  Mood  kejtt  at  a  temperjitii 
of  alioiit  20  ( '.  This  time  may  lie  eonsideralily  shortened  Iiv  iiicre.isi 
the  extent  of  foi'ciu'ii  material  with  whicdi  the  lilood  comes  into  <'onta 
iind  more  iiarticularlv    liv    whipjiinv:  the  Mood  with  a  iiuiicli  of  tvvij;s 


I'.l.diHi   (  l.oTTINti 


101 


; 


n\vrs.  In  this  liilliT  ciisc,  liowcvcr,  llic  clut  docs  not  rniiii  in  llic  usu;il 
Mi.innf?-.  lull  till'  line  tlncids  ol'  lilnin  ci. licet  on  tlic  twins  "f  wiirs,  Iciv- 
III..'  lifliind  the  IiIoimI  mtmiii  witli  tlic  (•oi-iiusclcs  still  sns|)ciicl<'<l  in  it. 
iiic  tiliiin  rcMiovi'd  in  tins  wiiy  ni!i.\-  liifii  lie  wjislwd  I'lci'  of  inlliciciit 
s,  iinii.  'I'lic  scrum  iind  iMnpnsclcs  nnw  I'mni  ih lihriinili <l  hhiml,  wliicli 
is  used  for  ni;in>  iiliysioloiiic  imrposcs.  Clottinu;  is  jdso  u'l'i'atly  acceler- 
ated liy  allow  ins.'  the  Mood  to  How  over  exposed  tissues.  Sonicthini.'  is 
.\ii|i.nlly  adilcd  to  it  I'roni  the  tissues  which  accelerates  the  clottinir 
j.roccss,  this   inlluenee   \><-\t\v  particularly    marked    in    the   ease   ol'   Mood 

,,l'  the  lower  vertelirates.     When  the  M 1  of  the  hii.l,   foi'  example,  is 

nreiveil  tlii-ousrh  a  cMMiiula  inserted  diieetly  into  a  \  essel  with  as  little 
iiijurv  to  the  walls  as  possible,  it  very  slowly  clots  if  at  all.  lint  soon 
does  so  if  the  Mood  is  allowed  to  come  into  contact  with  exccuiated 
lissucs,  iiy  if  it  is  mixed  with  tissue  extract,  smdi  as  that  of  muscle. 
Clottins;  is  consideralily  accel. 'rated  hy  warminu'  the  Mooil.  The  ap- 
plication (d'  a  (doth  or  tampon  well  wiiuil'  out  with  hot  physiolou'ical 
saline  to  a  wouiide<i  surface  is  a  most  cflicieut  means  of  allayin!.'  hi'm- 
nrrliaue   from   \cssels  too  small   to   liLrate. 


iil 


The  Nature  of  the  Clotting  Process 

riasnia  olitained  hy  centrifn<.'inir  hlood  that  has  lieen  prevented  from 
•  lottiiifT  liy  oiii'  id'  the  forcyoiny:  methods  can  lie  made  to  idot  li\'  rcmovinir 

the  inhihitint;  inllu •(■ ;  for  <'Xaiiiple.  in  cooled  plasma  iiy  warminu'  the 

Mnod  to  room  temperature,  in  salted  jilasma  hy  dilutini;-  it  with  a(   least 
all  equal  Volume  of  water,  and  in  deealcitied  plasma  hy  adiliiiir  a  siitVi- 

.ii  lit    ; Hint    of   soluMe   calcium   salts   to   conihine    with    all    the    added 

ii\alate  and  leave  a  snudl  trace  of  cilcium  salts  in  excess. 

'I'lie  (irst  (piestioii  coiiccrns  tin  sDiiin  uf  lli<  jihrin.  and  the  a'lswer  to 
it  is  furnished  1).\  eomparitiir  the  composition  of  hood  plasma  with  that 
"f  seiUTU.     Tholliih   hoth   of   these   fluiils   contain    the    prote  ns.    alluimin 

and  tdoliulin,  ill  approxiiiiat(dy  the  sai iicentratiiMi>-.  the  plasma  also 

i-.iiitains  niiothcr  ])rotein  not  unlike  ■iloluiliii  in  most  id"  its  reactions, 
I'lit  disliiiiiuishcd  from  typical  irlohulin  in  that  it  is  iMi'cipitated  l>y 
■lalf  saturation  with  sodium  (diloride,  ,iii  whiidi  typical  ■.doliiilin  is  solu- 
lle,  aiul  is  more  readily  coa<iiilated  liy  heat.  To  produce  half-saturation 
I  the  jilasma  with  sodium  (diloride,  eipial  volumes  of  plasma  and  satu- 
ited  sodium  (diloride  solution  ai-  mixed  together.  The  i>re<dpitate  of 
•hriiiinii  II,  as  the  sulistancc  is  called,  is  then  collected  at  the  liottom  ot 
'ii«  tiihe  hy  centrifiii-'ini;  and  is  washed  several  times  liy  d  eaiitation  with 
1  all  sjiturated  s()diuiil-(dil(iri(le  solution.  The  washed  precipitate,  dis- 
-'Ked  ill  weak  saline  solution  ( iir(d'eralil.\  containiii!:  a  trace  of  Mear- 
i'  iiiatei,  will  then  he  found  to  (dot   under  certain  conditions. 


U...J^J 


1ICJ 


mi:  c.i.nuii  AM>    iiii:  i.VMi'ii 


'I'lic   li('\t    i|il"stiiipi  (■(iiicii'iis   llii    iiiiliin    III    llii    i-iiihlil lulls   tliiit   iiiiisi    llii 
jihriiiiii/i  II  In  ihil,     Wlii'ii  ;i  liliriiinjtii  Mihilinn  is  rnisi'il  \  itli  a  I'cu   (Ir'nps 

111'  M I  sii  Mill.  ,'i  rill)   UsiimIIv    In  nils,  w  III  I'll  liipu  ex  cr  i>  int   I  he  ciisr  w  licii 

|)hi^iii:i  is  :i(|ilci|  iir  w  lien  llif  sciiiiii  is  licjiti'il  Ih'Iciic  .-Mldiii!,'  it.  Uci-iilisc 
:i  siiKill  i|UMnlil\  111'  sclllln  is  c-ipjllili'  nl'  (■;ii|siii'_'  tllc  clnl  "  illir  "f  ;i  liilui- 
ijli'llll  il.''  'iT  tilililliiu'i'li  -nllltioll  ul-  |pl:isMia,  il  is  sllppusnl  lull  Illl-  i|c|i\' 
slllist,iiii-c  |iii'sciil  ill  il  is  (iT  llic  ii.-ltlllT  111'  ,'l  riMiiii'lil  lihriii  h  run  III  nl 
tliriiiiihiii.  It  litll^l  111'  |iniiitni|  niit.  In  i\\r\  el'.  tli;it  lliric  i-.  I'linsiiliTJilili 
ilmilil    \\lirlli(i-  lliis  ai-li\i'  liiii|\    Is  ii^Mil.\    111'  llic   niiliiri'  nl'  m    rfrniriit    m 

I'll/Nlinv  i-'nl-  r\;illl|ilr,  ,1  It  In  ill  i:  Il  IliMtlil  si-llllll  lines  lint  cMjs,.  <-|llllilr^, 
llllntlllilll.  |ll  l|i,l|li|  rinlll  si-lllhl  ll\  llln  lll.'llliiil  .llinlll  tn  Im  i  Ifsl'I'il  in  I,  ill 
lllr  mIisi'IIiv  111'  iliii|-v;ilni'  s.llts  nllll  \i  it  llst.llnl  i-\n||  il  linililin  t'lll|icriltlllr 
MnlnnN  IT,    true    rii/v  mi's    iirr    i- li;i  lllrl  r  li /ri  I     1 1,\     tllr     I'.lc-t     tll.'lt,     liki'    lillin 

i';iIm1\  til-  .•I'ji'iits.  :i  \ci\    miniiti'  i|ii,intily  r.iii  i  I'l'ii-t  ;[  i-li.iiii;i'  in  :iii  imli'i' 

illill'  llllliillllt  111'  slllixliill,-,.  witllnlll  llic  lll/\li|c  liiM-nlnilln  llsril  |||)  ill  tin 
priil-n^s  I  |i'|'jr  7"_'  '  .  will  II  t;lln!l|lin  is  ;il|n\M.!  In  net  ll|inll  :\  li  I  ll'i  I  ln'_:r  1 ! 
snlutinll.  nil  tlif  ntliil'  |i,ll|.|.  it  i^  -,-|ii|  lli^it  nlllv  :|  lixi'il  llll|n||||t  n  I'  lilllih 
IMII  111'  rnlllll'il  W  lli'll  ,1  ^111,1  I  I  :illlnl|llt  nl'  t  II  M  ml  li  1 1  is  il  ill  It'll.  \l' it  1 1 1' I'  l|lli'~ 
this    ,'lllliiUll!     illi-1  I'.'l^i'    \\  lli'll    till'    tilili'    nl'    I'l'iii't  inn     is    |llii|nM'_'i'i|. 

\Vli;ili'\  I'f     lll,'l\      111'     till'    sii_rnili.'illiri'    111'    till'     rnlCMilill'^      r.'H'ts.     it     is     illlpnl 

tillit     tn    l,l|ii\S     lll.'lt     ll Ilitlill'J    suliv|i||ii-r',    I  lll'iilllllill,    r;in     111'    i--nl;itl'l|     I'lnlll 

lilniiil     vi'iiiiil     ill     .'I     liili'l  .'llil  \       pnii'     riillilil  ii'll.         I''nl'     this     plll'pnsi'     111 I 

SCIIIIII  Is  .'illnwi'il  tn  st.'lllil  limll'l'  .'1  l.'imi'  Xnlllllll'  111'  ;ili'ii|lii|  fill'  ;|  WCi'k  n' 
tWn;    till'    pi ipil.'ltc    i--    thi'l Ill'i'tiil    ;il|i|     nllilll'il    Up    witll    WJltlT,    wllii'i 

i'\li'.'ii-ts  tlic  lliiniiiliin  ii'niii  it.  Ic'uiiii;  till'  sci'iiiii  prntcin  in  ii  (•nimul.'iti  ■' 

sliitr  'I'lii'  li'Nllltinv  W.'lli'I'V  snllltinl!  Ill'  tlllnlllliill  lllilV  he  rilfllll'f  JH' 
I'ipit.'ltril     liy    ,'l|. -1111111.     till'     pl'li-ipit,-|li-    W.-lslli'il     ill     ;ili-nllill     .'lllll     ll'ljissiiKr.! 

ill  Wilt. -I',  \icliliiii;  iilliiii.-iti'ly  .-1  siiliilinii  \\liii-li  cxliiliits  \  ci-y  m.-irkcl  i-i. 
.-iirnl.-il  iii'j  piiHi'i's  hIii'ii  ,'|i|i1ii|  tn  phisiiiii  ni'  liliriiiiiL'i'ii  snliitinn.  TlirniM 
liin  slinws  must  111'  till'  pi'iitciii  ii',-ii-t  inns  Imt   it   is  imt  i-n,-iu\iliitcil  li\'  lici- 

As     wnlllil     lie     c\pi'i-ti'i|.     ,'|     i-nlisiili'l'iililc     ipMllt'ty     nf     t  II  I'nilll  li  II      |-rliniil- 

.-nllici'i'iit  111  till'  tili|-iii  rni'iiii'il  in  tlic  pini-css  nf  (-Initinu'.  iiinl  Ilnwc!' 
ili'si-rilii's  ,'i  \.'r\  iisi'lnl  imcIIiihI  In  wliii-li  it  ciii  lie  si-p.-initcil  fi'nin  lilui 
.-mil  pi-csi  r\  I'll  in  a  ilry  i-niiiiilinii.  itriclly  stalnl,  this  ini'thnil  i-nnsi^' 
in  all-iwint:  washcil  I'liiin  tn  staml  nxi'iiii^ht  iiinlci-  ciylit  jici-  i-i'  ' 
snililllii  i-hlniiilc  snlutinll.  wliirli  ilissnhcs  the  t  ll  I'nill  I  li  II .  The  l'eslll|i||. 
extract    is   then    nii\ci|    \\itli    an    ci|iial    Mililliie   nf   iii-elmie,    whieli    tliinw 

ilnwn   a    pl'eeipitiit ililailiilin    the   thiniiiliili.      Tn   p|-esci'\e    it,   the    prei-i| 

ilate   is  i-nlli'1-ti'il   nil   a    iiuiiiln'i'  nf  small    tiller    pjipei-s.    wliieh   ;ii-e   siili^i 
.plcliti>    npcjH  ij  i.iil   .III. I  liiic.j   ii\    cNpiisiiic  In  a   eui'lclii    nl'  enlil   air  ln-f" 
an  eleetrie  f.'iii.      When  the  thiniiiliiii  snluliniis  ai'C  ijesit'ctl,  tllc  dried  J)i  ■ 
eipitates  are   cNtiai-li-d   with    a    little    water. 


III.DUli    l'l,l>T'l'IN(i 


\m 


Tlironildn  tlocs  not  exist   in  lilond  pliisiiiii.  for  il'  ii  rlciiii  iiiiil  imiiifliliril 
..'liiss  tiiltc  is  iiisi'ilcil  iiitn  iiti  iirlcry  iiiiil  tlic  liliiml  colliTtcil  imdcr  ill 
idlidl,  llic  pici-ipitMlc  iit'lcr  sliimliiiir  a  IVu   w.'cUs  will  yield  int  tlii'oiiilpin 
uheii   triliiiiiled   «itli   wjiter.     <jiiili'  clearly,  tlierefcire.   tlie  llin.inl>iM   is 
|,iiidiired  Ml    llie  lime  tlie  hloiid  eliils,  and  llie  i|nesliiiii   arises.   Wtiat    is 

II  |.riidiieed  i'n>ni  .'     It   will   I'l-  r.Miieiul>ereil  tliai.  wlu'n  tlie  lilimd  is  ex 
;iiiiiiied    nnder    the    niielosecpe    dil-ilit:    the    el(iti;ii'/    |>nieess,    tlie    (ilinii 
Ihreads  are  seen  to  start   frimi   foci  uliieli  enrrespipiid  Id  tlie  lilticd   plate- 
lets.    It  wiiidd  appear  tlierel'ore  tliat   the  thrdliilpin  liilisl   lie  derived   iVuiii 

Miiiie  siihslal that    is  shed    r.>rlli    Ircim    the   platelets   diiiinir   the    disiii- 

le..'raliciii  which  they  iindery:<i  sIkmiIn    aftir  the  lil I  is  shed.     Tin'  siili- 

s|;iiicc  is  called  iinilliniiiihiii.  The  platelets  i.r  their  pieciirsnrs,  the 
Tiiei:acar\(ic,\  tes  oT   red    hniie   marrow,   are   pmlialil.N    mil    its  olilx    soiircc. 

t'cir   ehittinu    liia.N     i nr    iir    the    emiiplete    ;il>siMii-e    (if    platelets,    when    it 

.ippears  I.)  conn'   rnnii   the   lein ytes.      I'rol lirmiiKiii   appears   pleiitiliilly 

III  the  Ihiid  nsi'ij  In  perl'iise  reil  hniie  niarn.w  mitsiile  the  hi.dy  i  Drinker 
;ind   |)rinker''). 

To  Sinn  up  what  we  have  so  t'nr  learned,  it  may  he  stated  that  the 
process  of  clottini;  starts  with  the  disiiitetrral ion  of  Mood  platelets  and 
piohalilv  of  leucocytes,  as  a  result  of  which  there  is  shed  forth  into  the 
plasma   a    suhstance   called    prothrondiin.    which    imme.liately    aflerwanl 

I nines  activated   or  verted   into   thromhiii.     The  thromliin   then   at 

tacks  a  protein  present  in  jdasiiia  called  lihrliiii<i,  ii,  piodilciii!;  from  it  in 
tliicad  like  form  the  iiisoliil.lc  protein,  lil.rin.  Unt  this  does  not  com- 
plete  the   history,    for   at    least    two   other   important  factors   come    into 

play:  tin e  is  the  presence  of  soliihle  calcium  salts,  and  the  otiier  that 

of  peculiar  silhstanees  dcrivi'd  from  the  tissues  outside  the  Mood  vessels 
and  called  tliroinhoplastii  siilistances  or  tliromlioplast ill  (  Howell  i.  We 
must   now  consider  fin   ,.clioii  of  these  two  factors. 

The  Influence  of  Calcium  Salts.     As  already  explained,  the  i)roof  that 

-nJuMe  .••ilcinin  salts  aie  necessary   for  clottinu'  is   furnished   hy   the  oh- 

•■i\atioii  that  the  process  is  entirely  ]ire\cnted  when  the  freshly  drawn 

iloud  is  mixed  with  soliihle  oxalate.     To  this  iiroof,  liowever.  oh.icctioii 

mr-'hl  he  iiiadi'  oil  tlie  score  that  the  oxalate  /» /•  .s(    inhiliited  the  clottini:. 

That  such  is  not  the  case  is  indicated  hy  the   fact   that,  if  the  oxalatcd 

1  ^.nd  or  plasma  is  dialyzed  a-raiiist    physioloj:ie   saline  solution   till   all 

r  s(,liihle  oxalate  has  hceii  removed  from  it,  clottinu  is  still  ahseiit  hut 

.11, mediately   sniierveiies   if  some  soluble   calcium   salts   i.re   added.      The 

1.  icstimi  arises  as  to  how  tlie  calcium  ion  acts.     Two  possihilities  exist: 

i     that   it   is  concerned  ill  the  conversion  of  ilhrimej'e!!   to  lihrin.  n.iid 

-     that  it   is  necessary  for  eonvertiii'^  prothroiiihin   into  Ihromhin.     Il 

all  ipiilc  ii'adily  he  shown   that   it   is  l>y   the  second   of  these   i)roecsses 


if-    . 


f    f? 


i 


J 


nil 


Tin;  iii.iMHi  AMI  Tin:  i.vmi'II 


tliiit  the  (■.ilciiiiii  iicis:  I'di-  cxjiiniili',  cloltiiii;  (icciirs  wlicii  piliilicil  lln-niii 
liiii  is  iiil<l('<l  111  (li.ilv/cil  iixjilati-  IiIiiimI  m-  pl.iMiiii  iii-  to  n  pure  solution  oi 
(iliriiiii!,'i'ii.     Citriitrs  picvciil  cloitiii;:  liy  I'liiiiiini.'  cjilciiiiu  citratf.  wliirl 
iiltliiiii«:li  Miliililf  (lues  not  ionize  in  solution.     .,   is  tlit    I'rcc  i-Mlciiiin  ion 
liiiit  nil-  iinpoitiint.     The  iii-tioii  ol   tlic  itinniilc  is  soincwliiit   iiivstcriou-., 
for  it  li;is  liiin  I'liuml  tliiit  to  piodin'c  clottinir  in  (luoiiilc  plasma  the  siiii 
|>lf  adilition  of  calciinn  i-liloiii|i'  «ili  not  sutlicc;  tliroinliin  itself  must  I" 
ailileil  as  well.     Some  aniliois  assert.  Imwever,  that    if  the  ealc-ium  eliln 
riile    is    ailileil    eantiously    to    "tluoiide"'    Mooil,    it    will    imluee    elottiiiv 
I  Kiltu'ei-'.       In  any  ease  it  appears  that  the  lluoiiile  iloes  soniethiui,'  mon 
than   iii-ecipitate  the  ealeiuiii;    possibly    it    pievents   the   lireakiii^'   up   >>\ 
platelets   ailil    leueoeytes. 

The  Influence  of  the  Tissues.     .\s  alr.-aily  state.I,  when  slowly  elotfint: 
MoikI,  like  that  of  a  hinl.  is  eollecteil  thiotiu'h  a  sterile  «lass  tulie  into  ,i 
tliorou-rlily   elean    vessel    and    imim'diately    eentrilii'red.    the    plasma    wil! 
often   remain   indelinitely  iinelotted.      If  an  extraet   of  some  tissue,  siiel 
as  muscle,  is  added,  however,  the  plasma  immediately  (dots.     To  a  mud 
less  deH:rei'.  the   same   plienomeiion   is  cxhiliited   hy   manniialian    plasm, 
when   it   is  eolleeted  in  a  similar  manner.     From  these  observations  tin 
eoiudusioiis   may  he  drawn   that    the  tissues   furnish  souu'  sulistaneo  a- 
sistinu'   ill   the  elotliiiy:  looeess.   and   that    this  sulistaiiee   is  also   form.   1 
from   ei'itain  elements  lueseiit    in   mammalian   Imt    not   (iresent   in   avia 
Mood.      The    ah-eiiee    of   platelets    from    the    latter    Iilood    sujru'ests    th.:' 
tliese  must  lie  the  soiiree  of  the  aelivaliiiu'  sulistanee  in  mammalian  IiIium! 
Ft  is  plain  that  this  tissue  factor  in  (dottinj;  is  of  importance  in  liastc 
iiiLT  the  |)ioeess  when  an  animal   is  wounded. 

liefore  attempting  to  formulate  an  hypothesis  ihat  will  explain  tl 
process  of  elottiii}.'.  it  is  necessary  to  call  attention  to  one  other  imp" 
taut    fact.     This  ret'ers  to  the  lueseiice  in  lilood  of  a  sulistanee  that   pi' 

vents  elottiii'r  and  is  hence  called  (nitilhriiiii-lilii.      Antithi Iiin    is   pr. 

cut    in   normal   hi 1,    for  a    iriveii   specimen   of  ]iure  lilirinofreii    will   cl 

less  rapidl\   when  mi>;ed  with  serum  to  which  some  oxalated  jilasiiia  h; 

I II   aiMcd   than   with   an  eipial  amount  of  the  same  serum  eorrespoh 

iiit'ly  diluted  w'th  a  solution  of  solulile  oxalate.  A  strikintr  iiicrci 
ill  the  concentration  of  aiitithiomliin  in  lilooil  can  he  liiouixht  alimit  I 
rapidl,\-  iiijectiii!,'  a  solution  of  commercial  jieptoiie  into  the  Mood  \' 
sels  fifteen  to  Thirty  minutes  licfii:e  lili.edin-,'.  The  |iciitoiiized  lilood 
Iilasma  will  remain  fluid  for  many  lioiirs,  if  not  indefinitely.  T!;at  t' 
failure  of  tliis  Mood  to  clot  depends  on  the  presence  of  some  antielotti 
Kuhstance,  and  not  upon  the  ahs.-ucr  of  one  of  the  neccssai-y  clofli;.,-  s:. 

stances    ( tilirinou'cn,   tin iliiii,   etc.  i.    is   evidenced    liy   the    fact    that    it 

.•iddition  of  .some  of  it   lo  a  mixture  of  throudiin  and  fil)riiioj;en  inhii' 


III.<M)I>   »I,()TTI\.) 


i()r> 


the  (Miiifrulali'iii,  wliicli  it  (li»s  imt  do,  luiw ever,  it  it  is  liist  ol"  till  licntfil 
to  SO'  LV  mill  liltficd  I'liM'  lit'  tlic  (•initriiliitfd  pintcin.  .Morciivi-r,  tlu' 
.iiitiifriiiiistic  fii'tiiiii  is  i|ii!iMtitiiti\i'  in  llic  sfiisc  tlint  a  liscd  aiiiotint  of 
the  |)('|)toii('-|)liisMia  iidiiliits  tlic  action  of  a  ti\<'d  amount  of  tlirninliin. 
The  soiiiTc  of  antitlii'oniliin  in  tlic  l)ody  appears  to  lie  mainly  at  least 
ilic  liver,  for  it  lias  hccn  found:  (  I  >  that  peptone  injection  into  an  animal 
iVoiii  wliicli  the  liver  lias  licen  removed  iloes  not  ea\ise  aiitithromliin  to 
lie  formed  (Deniiey  and  Minot  )  ;'"  CJ  i  that  peptone  injections  into  the 
poital  \ein  cause  antithniiiliin  to  appear  in  the  hlood  niueli  nioi'e  rap- 
i.jly  than  when  the  injection  is  made  into  a  systemic  vessel;  and  i^Si  that, 
when  the  liver  is  perfused  outside  the  liody  with  a  perfusion  lliiid  eon- 
laminir  lu'ptoiie,  antithromhin   aceiiiHulates  in   the  i)erfusion   fluid. 

.\  (luid  contaiiiinjr  a  liijih  eoiicentration  of  aiitithromliin  is  secreted 
li\  the  so-called  salivary  jrhiiid  at  tlw  head  eiicl  of  the  leech.  Tlie  func- 
tion of  the  Iliiid  is  to  i)revent  clottiiiir  of  the  li|oc«l,  so  that  the  animal 
may  continue  to  suck  it  without  interference  hy  elottintr.  After  apjily- 
iiii:  leeches  for  medicinal  juirposes  it  is  thercfoi'e  necessary  to  wash  the 
wound  thoroiiu'hly  with  water  so  that  all  traces  of  the  aiitithroinhiii  may 
III-  ri'moved  ;  otherwise  the  hleedin^'  ma.v  continue  for  a  coiisiderahle  time. 
Practical  use  is  iiuide  of  this  ctTect  of  the  leei'li  to  ))revciit  clotting'  of  Mood 
outside  the  liody,  or  it  may  he  used  to  inhiliit  coiiriilatioM  lulni  rilaiii  in 
experiments  wheiv  elottinsr  would  otherwise  interfere  with  their  ])r(if;- 
ivss;  for  example,  in  crossed  circulation  t  xpcrimeiits  (  pajre  oti.'i)  ami  in 
i'N|M'iiiiieiits  in  vividitVusimi  fpasre  (KIT).  For  such  purposes  the  leei'li 
Ihad  is  cut  off  and  extracted  either  with  saline  or  iiy  treatment  with 
I'liloroform.  which  removes  other  jiroteins  from  the  saline  solution  leav- 
iiisj  a  stronir  aiitithromliin,  known  under  the  trade  name  of  "hirudin." 
At  temperatures  about  that  of  the  liody  the  action  of  aiitithromliin  is 
-iie.itly  auv'iiiented.  In  animals  like  the  inatmiials  in  wliidi  the  content 
'<r  anlithroiiiliin  is  small,  this  may  lie  important  in  mainlainiu'.'  the  tlu- 
I'lity  of  the  Mood  (Howell).  Itlood  containini;  aiitithromliin  can  lie 
•adc  tci  clot  liv  the  addition  <if  throniliin,  and  therefore  of  lilood  serum. 


'<i  jt  4'^  I 


t  IIAITKi;   Mil 
I'.l.otiji    Itl.uuji  r|.o'|"|'|\<;  irciMt'.li 

THEORIES  OF  BLOOD  CLOTTING 

.\lliiii|ils  III  link  ^ill  till'  riirryniMir  I'lu-ts  |ii!.'rllicr  ill  \\\f  I'cuiii  «(  i 
silillilc  lliriii\  li:i\i'  Mill  ^11  {'-.w  lii'i'li  cliliiciv  MiiTcvsrilJ.  All  .liilri'  lli:i' 
till'  liliiiii  is  ili'i  ivi'il  rimn  (iliiiiiii;;i'ii  |i\   till'  iirliiiii  nl'  tliiiunliiii.  tlir  |Miiiii  - 

ill    ill>|illtc    liriiiii    tliiisi'    wliii'l lii'cili    the    (ili'.'iii    lit'    liir    thlutiiliill    jiimI 

llii'   liiiiilr   lit'  .irtiiili    III'  liii'   (•.liciiiiii   iiliil    tliriiliiliii|iliislir   slilist.iliiTS,      'I'li. 

tlniiiv    iiiiist    wii|i'l\    ;iiTi'|(lii|    ill    i;iiiii|ir    i>   lli.il    111'    Miiniw  it/,,   ,• iiliii 

In  \^llil■ll  till'  llii'iiiiiliiii  i'\i>ls  ill  li\iim   liliiiiil   ill  nil    iii;irti\c  sljiti'  ci'lli' I 
till  iiliiliiiUi'll    I  |il'iitliliiliiliili  I ,    «jiirli    lirciiliirs   n  iini'il  ri  I    illtii    I  liri'lllliili    li'. 

till'   siiiiiill;ii IIS   :icliiiii   nil    it    111'  siijiiliii'   i-iili-iiiiii    siills   Mini    nf   tliriiiiili" 

I'l.islii'    siilist.iiii'i  s    I'm  llislicil     liy    till'    lisslli'    Cfll.s    ill    jiclirlill    illlil    li,\     \\i- 
rclh(l,-|r    1  Irllinils    nl'    lllr     lljnnil     jlhllclcts    :illl|     lcl|(Mlc\  tcs.        Al'CllI'llilli;     |., 

Iliis  \ii'\\    till'  I  lirniiiliii|ihislii'  siilistiii ,  iiiili'il  liy  tlic  picscin t'  ciiciiin 

inlls.  rnliM'lls  I  ll  rnll  In  liii'li  <  |  lint  ll  I'nliilii  I|  •  til  t  ll  rnlllllill.  It  ill-Is  tlliTr 
I'liir   ,is   ,1    LiiKisr   .iliil    is  cilh'il    tllldllllinkin.isi'.      '{'he    t'lllliliinii-ntill    nirl    nl 

lliis    llirniv,    Ijii'ii,    is    lliiil     l,i|i;isi'    is    1 ssiiiv    I'm-    till'    Ullinli    {<{    till'    ra  I 

iMlllii  with  |iriitliriiliiliill  M  l';icl.  lin\w\c|-,  wliii-li  is  rliiill('lii.'r(i  liy  lliiWiH 
\\  III!  sl.ili'S  tllllt  liriitlilnliiiiiii  iii;iy  111'  rnlix  crtnl  In  t  |i  rmilliill  l)\-  tin-  iii-lin 
nl'  ralriiiiii  imis  aliiiii'.  'I'liis  iii\  cstitxalnr  liclicx cs  lliiil  tlic  tliiniiil.M 
plastic   siilistaiii-i'   ai-ts   iiiil    as  a    kinase    liiit    liccaiisi'    it    iiciiti-ali/i's  an' 

tliriiniliin,  wliii-li   is  i'iiiislaiitl.\    iHisi'iit    in  tli<'  M I.  ami  tlii'  fiiiictinii 

wliirll    is    tn    |iii'\rn|     l| ali-illlll    I'inlll    llliitiliy    \\illi    till'    pi'ntlil'iiliiliin    ' 

I'lii-in    tliiniiiliiii.      llnwiU's    li.,iiiy    in    liis   nun    wnnis   is   as    t'nllnws:  "I 

till'     l-il'l'lllatinu'    liliinil     \\v     lillil     as    rnllslant     I'nIlstitUCIlts     lill|-ill(ll.'t'M,     pi' 

tliiniiiliiii.  i-ali-iiiiii  sails  ainl  a^il  it  hiniiiliin.  Tin'  last  iiaiin'il  sulistaii'-' 
linlils  till'  priitliiniiiliin  in  I'lmiliiiial  inn  ami  lliiis  pri'vciits  its  ruin  iTsi' 
111-  arli\atiiiii  tn  I liriiiiiliin.  Wlicn  llir  lilmnl  is  slicil.  till'  (lisiiitcirrali- 
lit'  till'  I'lirpiisi'li's  I  plairli'ts  1  I'uinislirs  iiiali'iiaj  i  llimiiiliDplasf  in  )  wlii' 
I'liiiiliiiirs  with  111!'  antilliiiiiiiliin  ami'"  at  tlii'  same  tiiiic  lilici'iitcs  iii" 
"  prnlliiniiiliiii;  tlir  lattiT  is  tlirii  ai'tivalril  liy  the  calciiliii  iiiiil  arts  ■ 
the  (ilii'imiL'(-n."     Ant  itliinniliin  can  alsn  prevent   the  action  of  llirmiil' 

nit     (ilil-illniT.r,.        A-    alrc^i'ly     pnjiitc.l     ..Ijt,    tliv     t  l.rtrlhlMijlla.st  ill     cat!     lie    ■': 

ii\eil   t'lniii   tlie  III I   itseir  ill   the   iiiaiiiiiials,   but   only   fnnii   the  tissii 

ill  the  Inuer  \  eileliiales.     It  is  iiileiest  iiiir  tn  iiiite  that  the  thliiinliiipla^' 

liii; 


iii,(MM>  rruiriNc 


107 


,111  III'  l'\tl'il<'li'<l  llnlll  llli*  lisMII's  li,\  lilt  soKi'llts.  ami  tllllt  it  M|i|li'ills  tn 
iiliinu;  til  till'  ■lavs  nl'  |l|iiN|i|iilliiN.  Iii'illl.'  ilnliril  rliisriy  Iclatnl  to,  if 
<>l    iilrlltiral    \\itli.   krlilialili   lllnwrlli. 


Intnivoiicular  Clottin<^ 

'I'lir   |itaiMiral   a|i|ilii-al  imi  nl'  llir  llii'iirv    nl'  IiJikmI   ilolliin.'  i-hihtiiw  tlir 

riiaiinrr  in  wliirli  tin'  MimhI  is  iiiainlaiiii'il  in  a  lliiiil  riiinlitinii  in  tlir  M I 

\i>M'is,  anil   till'  dislml'ani'i'  nt'  llii>   I'liiirlinii   raiiNiii'.'  iidnicii'iriihir  rlnl 

tnhi.     Ai'i'iii-iliiiu:  tn  tl Ill'  tlii'iir.v,  till'  III I   is  inaiiitainril   lluiil   li\    tin' 

alisinrt'   rriilii    it   III'  an,\    cnnsiij)  ralili'  i|naMtil.v    nl'   Uinasr,   ami   a nlitiu' 

111  till'  ntliiT.  li.\    till'   |iii"«<'m-i'   in    it    nl'  an   aiiiniint    nl"  antillirniiiliin   sulli 

rlilll     tn    jiri'M'Ilt     till'     llllinll     .il'    rall'iuill     willl     |llnl  ll  l-nllll'i  11 .        Till'     lllliilitV 

I-  iiiainlaiiii'il  cmii   wlnn  lai'ji'  anmniits  nl'  tlnninliin  nr  nl'  lilnml  siiiiiii, 
\\lii.'li  cnntains  this  Mil>stam'r.  air  iiijii'li'il   iiiln  llif   liviiiu:  animal.      Wf 

r;lll    lust    CSIllain    till'    il unity   nl'    lllr    III 1    In    till'    arlinli    n  I'    llllnrilllin    nil- 

il.r  tlii'si-  cii'i'iinistani'rs  as  ln'iii'.'  iliii'  In  tin-  iiislanlanrniis  a|i|ii'arauii'  in  it 

■  ii  antitlirnniliin  in  aninnnts  snt'lii'ii'iit  tn  |ii-,'\i'nt  tin'  arlinn  nl'  tlirnniliin 
.I'l  tiliiiiiii'.'rii.  I'm-,  as  slalrii  almvi',  it  is  i-laiiin'il  li\  linwi'll  llial  anti 
ilii'niiiliin  lia'^  tins  inlhii'in'i'  as  wrll  as  tlial   nl'  pii'v  riiliiiy:  tlir  cnin  cisinn 

■  ><'  {'inilirniiiliin  intn  tlirmiiliin. 

I iiliiird.'ii-iiliir  ihilliiiii   iiia>    111'    liiniiulil    aliiiiil    I'v    a    \aiii't\'   nl'   nii'aiis: 

I      Mi'i-lianii-al  ilaiMai.'r  tn  the  lininu  nl'  llic  lilnml   xcssi'ls;  al'lrr  llir  ap 

I'lii'atimi  III'  a   liLralnri'.   rni-  r\aiii|ilr.   tin'  il.'Miia'j-i'il  iiiilntlu'liiini    is  snnii 

.•n\iii'i|  liy  a  rint,  uliii'li  '.'railiially  1 uiii's  liiini'i-  ami  liriiii'i'.  ami  iiiav 

>|iii'ail  ii|i  tin-   M'ssi'l   In  till'   iii'Nt    liianrii.      ■'-■    'I'lu'   in-i'si'in I'   fnii'iirn 

-ulislalirrs    ill    tin-    lilnnil.       Mlillinli,    I'nr    I'Na  lll|ili'.    aii'    apt     tn    railsi'    I'lnts 

'•'  I'niiii  at    till'   plai'i's   ulii'i-    tliry  stirk.   iiaiiirl,\.   in    tlir  sMialliT   vi'ssi'ls. 

'  '"Itiiii:  is  alsn  a  rinpii'iit  nrrniTcni'c  wlii'n  tlii'ic  air  Incal  ililatat  iniis  nl' 

1'    lanliiiv  asi'iiliii-  tiilii'.  ami  it   ina.v    m-i'iir  umli'i'  inipi'i  I'li'tlv    nnilrrstniid 

"I'lilimis   caiisinir  tlic   cnnilitiiiii    knnwn    as    I  hri  mi  In  is  is.      i'.\]     An    iiiti'i'- 

■'ii'_'  \aiit't\    111"  intiavasi'iiiai'  rlnttini:   is  tliat   faiisfil   liy   tin-   iiitia\«'- 

I-  iiiji'i'tinn  111'  salinr  rstracts  nl'  n'll  rii'li  tissiiis,  siii-li  as  tlic  tli\iiHls, 

iiipli     ylamls     nr    ti'stcs     l  WnnMl-iilui' ■ .        Uy     pl-i'ripitalinll     \\itll     aci'tii' 

'I  aihl  liiui'stinii   with   pi'plnin',  a    ii'siilui-  I'aii    lie  nlitaiiu'i!    I'nini   tlii'sc 

i';i'N  whirli,   uhcn  ilissnhi'il    in   alkali,   has  a    very    pinnnuni'cil    iiitra- 

'   iI'M'  I'lnttini;  I'lTi'i-t.     Since  tlii'sc  jiri'ripilati's  arr  \i'i>-  rirh   in   jilins- 

''.^,   it    is   prnlialili'   that    tlii'\    an'   nl'   tin'    natni'i-   nt"   phnsphnprnicin 

'  "iilliiiinin  I.     'riit'ir  actiiin   niiist   (Irprml   nn    ni'iili  ali/atimi   nl'  anti- 

il'iii,  ai'rnriiiii'_'  tn  llnwi'll's  tlii'ni'y,  nr  ln'caiisi'  thi'y  siTM'  as  ihrniii- 

^is   1  ari'iiniiiii;   tn  ^Iniaw  it/."    tlicinyi. 

a   Miattcr  nt"  fai't,  Iniwcvi'i',  tin'    ("niiL'nini;   iiIisi'i-\  alinn    is   imt    cnm- 
!•    ■■   >  rNplaiiii'il  ti\   citlicr  thcnry.     If  in  jilai-c  of  inakim^  mn'  injectinn 


Ills 


TIM;    III.IMM'    \M>    I'lll,    l.\  MI'll 


flc<|Uflit    ilijrcliinis   (if   Milllll    iiliinlMlts    of    tin-    iili"\c    liillli'i'iiil    lire    llliulr 

iiisli'iiil   of    iiitnixiisculiir   I'li.tliiiu',    a    ili-lny    in    tlic    iMiacnlatinii    li i 

liki'lv    ti>  oi-nir.     IikIi'ciI.  ri'pi'atiil  itijcriimis  nf  miuiII  aiiHMiiitH  iiia>'  n 
linly  rcrruivc  llic  rfolliiiK  poufr  nf  tlii'  \<\»i>,l.     The  ifadiiicNs  with  wliiil 
this  »ii>  failed  "iicualivc  pliasf"  appi'ais,  scctiis  tu  ijcpfiitl  mi  Ihc  niiln 

tixi lulilioii  of  tlir  aiiiinal  at  the  liini'  of  inji'i'tiiiM      If  a  larjrc  iIum.  i 

iiijcclcil  into  a  fastinir  <|ol',  fi>i'  (Aaiiiplr,  tlinmilxisis  is  ciiiilincd  tn  th. 
pnrfal  ana.  whcn^as  if  it  is  injcrtcil  iiitn  a  icfi'iitly  fi'd  aniinal.  tli' 
tlift.iiilMisis  is  iiiiivrrsal  tlironirliniit  tin-  vascular  systi-m.  Tin-  (li'Vclo|. 
inciif  of  Ihc  lu'u'ntivi'  phnM-  is  iiiHliiiilitcjIy  (Irpciulcnf   upon  somh-  ich 

tion  on  t/i-  Mart  of  flu-  Ii\  iiii;  crlls  of  llo-  orsjanisni,  since  it  does  not  ir 

oil  the  adiiilion  of  siniilaf  sulisfaiiees  to  Mood  outside  Hie  liody.  'I'li. 
reai'tion  is,  indeed,  akin  to  tliat  li,\  uhii'li  iiiiiiiiiiie  liodies  in  ireiieral  ai- 
produced.  For  exainph',  a  toxin  injected  in  larirc  amount  has  a  ci 
tain  toxic  cfTect,  hut  in  repealeil  small  doses  w'tli  interveiiiii','  intiTVal 
it  leails  to  the  itnitluetion  of  an  antitoxin.  So  with  the  siihstanoe  \< 
i|Uestioii:  a  lariri'  (hise  injected  at  one  time  causes  a  jtositive  cfTect  dot 
tinsr  hut  smalli'r  doses  freipiently  injected,  the  opposite  efrcet  want  <'^ 
elottini:.  It  i  pi'nhalde.  as  suifu'ested  liy  StaiTmir.  that  more  inteiisiw 
stuil.x  of  the  conditions  caiisini;  intravascular  elottiiiir  will  throw  cui 
sideralile  liulit   on  the  ireiieral  (|iiestion  of  the   production  of  immuiiitv 

Measurement  of  the  Clotting  Time 

To    mtiisiin     tin    rhitliiiii    linn    i,f   ilniini    siliiil>hs   nf   hlnnil,    several    cc.i; 
ditioiis   must    he   oliserved.      These   have    heen    tahulaled    l>y    Addis"    , 
follows: 

1.  The  specimens  of  Mood   must    always  he   olit.-iincd   hy   exactly   tl 
s;ime   tecliiii<'.      h   would   introduce  serious  errors   to  compare   the   cl..' 
tint.'   time   of  one  specimen   of  hlood    receixed    from   an    incision    of   li 
skin    tear   lolie  t    will,    that    of   another   c.pllected    in   a    syriii'-'c    li\-    vn 
puncture. 

2.  The  temperature  conditions  must  alwa\s  he  the  same.  I'lolial 
'ir*  ('.  is  the  best  temjieratuie  to  use.  lli'_'licr  temperatuii's  are  ummi 
ahle  for  two  reasons:  lirst,  hecaiise  diirinj,'  its  collection  the  hlood  w 
lia\<-  Imcoiiic  cooled  to  alH>ut  nr  Im-I.iu   this  point,  and  time  would  Im'  i-. 

sumed  in  raisiiii;  it  hiu'lier;  and  se id.  hecause  the  time  of  coaiiulati 

i''    niiire  and    i •>■   sliorleiied    for   each    <|ejrrce    that    the    tempcratur.- 

r;iised,    this   at leratioii    hecominjr   especiallv     pronoun 1    for    tem|i' 

tures  ahove  2."»     < '.     Quite  apart    fr the  liahility  to  incur  errors   in 

dl'Ut     tn    nieaslireliii.Mt    of    s!!"!te|-    j>eriods    nf    lime,    nh-.c!'.  :;!li>!i--    ;it     lii--'' 

teniperaturi's  necessitate  iimst   liyoious  ailherence  to  ;i  li.xed  ti   .  pciati 
of  the  w;iter-liatli.     Temperatures  mui-li   hcluu    'J.')     ( '.  are  unsuit.ihlc,  ! 


hr,iNir>  iLDTTiMi 


10!) 


ijuisf  III!'  rliilliiiu  Mts  ill  minlinilly  mill  it  is  (lilllnill  to  tell  |iiri-iHt't.v 
\>  htii  it  occurs. 

l.  Tlic  lilonil  iiiiiHt  iiluiiNs  I Heeled  in  the  same  sort  "f  vessel  ami 

...MIC  ill  eoiitiict  with  till'  saiiH'  kiinl  ami  aiiioinit  of  I'oreiKii  material. 
T.I  this  it  ma.\  he  a<lilcil  that  the  reeeivilitf  vessel  must  he  serupilloiisly 
.  Iciiii;  aiiv  trace  ol'  old  Mood  I'hit  or  of  serum  is  es|iceiall\  to  he  miarded 
.I'jMinsl. 

\  The  cud  point  must  lie  >h.ir|i.  It  is  heri'  that  the  irreatest  teidiliieal 
.hlliciilties  arc  met  with  in  malilii}.'  precise  measurements,  ami  it  is 
•/icativ  to  he  desired  that  ililTcieiit  invcstinators  shoiihl  ailopt  some  tllii- 
r..iMi  method.  l''or  e\pcrimciilal  purposes  the  method  of  ('aiiiioii  and 
M.  ii.jciih.'iii''    is   no   doiilit    the   hcst,   anil    it    has   the   added    advantage   of 

L'uinj;  a    ^'laphic    record    of    ll liM'rvali.ins.      The   aci paii,\iii^'    tinurc 

I'i'/.    10)    shows    the    priiwiple   of   the    melliod.      The    Mood    is    received 
i!iiiiilt;li  a  stalid.ird  cannula  (''  i   inio  a  lilhe  I  V  >  .">  em.  I'li-r  and  of  .'»  mm. 


TH 


T* 


If 


I  .1:     1'.      |....^.n.ii    .f    111.    »!T.i|.hi,    ,..,i^,.l..Tiu  1. 1.       I  h.     .:.i..iul.i    ..I    ll..     ti«lil    r.-t..l    11.    n    ».ii.r 
■'1    ii.l    -liiim.    Ill    ilii»    .h,i>!i..in.       r...     luiihii    .li~.ii|.l..ii    -I.     u\t.       ll'.....i    l'..i.n..n    aii'l    Mm 

h.ill  .• 

!i  inal   diamet.  '•     and   a    loop    lof  2    mm.   diameter)    at    the   end    of   a 

|.|M  r  wire  (/)),  wliiidi   is  S  cm.  loiijr  and  H.li  mm.   in  tliamcter,   is  al- 

Mil  to  fall  jrently  into  the  blood  at  regular  intervals.     The  upper  en<l 

till'  wire  is  articulated  with  the     hort  arm  of  a  liuht  lever  so  couiiter- 

MM(l  that  when  the  stop  ilx),  wiiidi  ordinarily  holds  it  in  a  liori/.ontal 

-ition,  is  released,  the  wire,  now  Iiaviio^'  a  net  weijiht  of  lit)  mir.,  falls 

the  hlooil  ill  the  tiilio.     The  loin,'  arm  of  the  lever  is  iirovidcd  with 

'  liiiiiij  point,  which   is  made  to  inscrilic  its  movements  on  a  dnmi. 

!.inu'  as  the  hlood  is  unelotted  the  loop  sinks  into  it    when  the  lever 

Iiascd  and  a  vertical   line   is  traced,   hut   whenever  elottiiiu'  occurs 

'.p  sticks  on  the  hlood  and  the  writin-;  point  iloes  not  rise. 

'  clinira!  jnirpuses  where  Mood  collected  in  a  syrinire  hy  venipuncture 

■  !.  the  method  of  Howell''  is  most  accurate.     4t  consists  in  placini: 


.!•   It     I'h.vii.il  ,    .May    I,    l'*14,    xxxiv,    No.    2. 


110 


illi:  hi.dun  wri  -iiir.  I.YMI'II 


■J  of   4   iM'.   (iT   I  III'    III I    ill    ;i    wide    till r  •_'!    llllll.    liiiiMlctiTl    tlliit    li:i- 

lici'ii  <'l(',iiii'l  liv  ,1  liicliiciiiiMli' Mciij  iiiixlnif.  Tile  in'riml  tli.it  cljipM^ 
lictxM'cii  llii  iiiiiiiii'iil  cif  tlic  ciitix  of  lliiiii  jiiti)  till'  syiiii'ic  and  that  :i' 
wliii-li  till'  rliit   lias  lii'riiiiic  linn  ciiiiiiifli  sn  tliat   tlii'  liilic  ran  lie  iiivcrtril 

\\itliimt  s|iiirnitr  any  lilnml.  is  laUcii  a>  tin'  I'liittinu'  tiiiir.     Siiiri'  tiic  lil ! 

iliM's  nut  rniiic  ill  iiiiitai-'  witli  I'Npiiscij  lisMii's,  it  laki's  truiii  "JO  to  ti<i 
iiiimiti's  til  I'liit   l>y  tliis  nu'tliiKl. 

I'°iii'  iiiiiliiii'  i-iinii-al  ■'saniinatimi  nl'  lilmiil  talvcn  Ir'niii  a  skin  wihiihI 
I'lriiilii'  ami  liiissrl's  iiii'l  liml "  is  iiinst  s.-itisjartiiiy.  Tiiis  I'unsists  in 
liriiii-ipli'  ill  iilisiTv  iiii;  a  iliiip  nl'  Mnml,  iiinli'i-  tlii'  low  |)ii\\('i'  iif  tli' 
iiiiiM-oM-Mpi',  xvliili'  a  liiii'  iMiiii'Mt  III'  air  is  •initly  lilnuii  au'aiiist  it  ;i' 
ri'irular  iiitiT\als  in  a  taii'^i'iitial  iliri'i-t  inn.  Tiitil  rlnttiiiLT  si'ts  in,  tin 
iiiili\  iiliial  i-nipiisi'|is  iiin\i'  I'li'rly  in  a  I'iri-iilar  ilii  ri'l  inn,  Init  as  sncm 
as  cliittiiii;  hi'iriiis  tlii'y  ninxr  in  iiiassi's  w  liii-li  si  nii  ti'lnl  tn  licroim"  lixnl 
sii  that.  allliniiL;li  lln'v  iiiom'  sminuliat  w  lirii  tlir  air  inipiii'ji's  nn  thi'iii, 
tlii'V  iniiiinlialrlv  iitnni  to  tlirir  miirinal  pnsitinn  w  lii'ii  tin'  cili'ii'iil 
is    ilisi'niiliiiiU'il.       Wlii'ii    I'Inttiiiu:    is    i'niii|)li'1i',    tlir    air    I'lii'i'i'iit     iiicnl} 


piTssrs  nil  till'  I'liipilM'Irs  al  nlli'  pnilit.  I'i>  this  iiii'tlinil  till'  rlnttili 
tiiiii'  a\i'ia'^i's  livr  iiiiiiiiti's.  A  rniivi'iiii'iit  apparatus  I'm-  this  iiictlmil  i~ 
llial  Ml'  l'..pi;i;s.  uliii-li  is  slmuii  in  \'\<j:.  I'n.  ||  i-nnsists  nl'  a  tl'iliiral<' 
rniii'  1)1'  '.^lass.  pinjri'tinu  intn  a  iimist  i-liaiiilirr  pin\  iilnl  with  a  liilii'  •' 
till'  siili'  sn  iiiiaii'_'i'i|  lliat  \\  lirii  air  is  Mnw  n  intn  tin'  rli.-iiiilii'r,  it  stiilv 
till'  ilrnp  III'  llllll. I    plarril   nil    till'  ciiil   nl'   till'  riiiir  t  alijii'lll  iaily. 


Blood  Clotting  in  Certain  Physiologic  Conditions 

I'.rsiilis  till'   iNpriiiiiriilal   I'liiiililiniis  alri'.-iil\    I'linnici'.'ilril   as  I'lianyin 

llti'    i-liilliii'j    liiiii'    in    till'    lilnnil    III'    laliiiraliir.\    animals,    s) ial    iinntin 

lllllst  III'  lil.'lili'  III  llii'  ilillllilirr  111'  I'piiii'phl'ilK'  iliji'i-tinlis,  iif  I'nllilitim 
siippiisi'ij  til  i-aiisr  a  li\  pi-i  sii-i  itimi  nl'  this  liiiriiiiinr.  «(  llic  I'liintinn 
ami    111'   Iniiiiiii  haul' 

I'^pilli'plirillr    Mililril    tn   ilr.iwii    liliiiiij    ijiii'S    lint    alVri't    lln'    I'liitlill!;   liiii 

liiit  it'  small  .•inmniils  aii'  inji'i-ti'il  iiitra\  I'lmnsly  nr  i'M'ii  snlinitani'misi 

a    niai'!'.'''!    ilc'ira^ I'lirs    .  t 'jiiin..!!    ainl    <i!ay:    el'.    < ':innnn,    Ini'.    i-i' 

{..'iru't'l'  injirtiniis  may  li;i\i'  tin'  nppnsiti'  i-lTi'i'l,  ami  iiiti'i'iiii'ijiati'  .■ininiin 


111. (M IK    Cl.dTTlNd 


in 


4^ 


may  ciiuso  at  fifsl  a  |)niliiiij.'a1iipii  anil  later  a  sliortfiiiii";  of  tlic  t'mi". 
'riicsc  I'csiilts  with  larsriT  dusi-s  air  rdati'd  to  IIkwcH's  olisi'ivatioii  that 
repeated  doses  of  relatively 
so  !,'rea 
pliilie. 


>e  aiiiimnts  of  epinepliriiie  in  doi;s  may 

tly   retard  eoajiulatioii  as  to  make  the  animals  praetieally  hemo- 

It  was  further  foiiixl  liy  Cannon  and  liis  eoworkeis  that  epiiieph- 

the   elotlin";   time    when    injected   into   animals 


line   does   not    inllnen<'e 

from  whii'h  the  alxloiiiinal  visei 

hv  lisration  of  the  inferior  vena  ( 


a  have  lieen  removed  from  the  eireulation 
•ava  and  the  ahdominal  aorta.     In  the  lij^ht 


if  the  intluenee  w 


hieh  destnietion  of  liver  eells  I  liy  phosphorus,  eldori 


form,  ete.)  is  known  to  have  in  lennthenin-r  elottinir  time,  it  would  seem 
r.asonaiile  to  eonelude  that  it  must  he  throU!.'h  this  ortran  that  epineph- 
rine develops  its  eloltiny:  elTeets. 

Stiiinilation   of  the  splaiiehnie   nerves  j.lso  shortens  the  elottin<r  time. 

:ind  it  would  ai)pear  that   tliis  aeti lepen.ls  on  the  resuliiu!:  hypi-rse 

cretion  of  .'piiiephrine  ( |ia^re  T4(ii.  for  it  is  not  observed  following'  stniiula- 


ti.m  of  the  nerves  in  anima 


Is  from  vvhieh  the  adrenal   -rlands  have  lieen 

le 


,\cised  (Cannon  and  Xlendetdiail ).  The  inteicstinjr  su'.'U'esI  ion  is  mad( 
liy  Cannon  that  the  shorter  elotlin<i  time  ohserve.l  in  animals  showinir 
slroiiii  emotions  of  fii-Jit  uv  tear  may  also  he  due  to  the  liyperseeretioii 
,,r  ..pinephrine  which  this  worker  lieliev.'s  a( iiipaiiies  such  states. 

Blood  Clotting  in  Disease 

With  the  fill-tors  coiieerned  in  the  process  so  w  ra|)ped  in  mystery,  it   is 

not  siirprisinsr  that  the  nnderlyinir  <'anses  responsilile  for  i|ela.\fd  or  de- 

rK'ient   clipttiiif;  of  Mood   in   diseased   conditions  or   for  the    lornialion   of 

inliav.-iscular  clots  (ihromliii   are  little  understood,      .\cconlini;  to  How- 

.  ITs  theory  »(  the  nature  of  the  proi-ess.  which  is  the  most  satisfai-tory  at 

ihi'    present    time,    ahnorniai    clottinji    mi-iht    he    due    to    the    follou  inu' 

•  Miiscs;     (  1  )    ,1    'llniiiiislixl  amiiiiiil   iil   jihr'niinii  ii.     This  occurs   when   the 

!iipa1ic  eells   are    frrcatly   dajnaireij.    as    in    poisoiiintr    hy    (  hloiofoiiu    or 

;'lio>plionis  and    in   such   diseases  as  acute   yellow    atrophy   and   yellow 

\'\fv.     In  man\   eases  of  chronic  cirrhosis  of  tin'  liver,  as  Whipple,  etc.,'' 

..\c  shown,  the  hlood  also  clots   fcchly   heciiise  of  dcliciciil    tilninoiren. 

!;   sl.tiiihi  lie  pointed  out   that   it   is  not   so  much  the  cloltinn  lime  that   is 

•icase<l  in  such  eases,  as  the   lirmne.ss  or  coiisistincy   of  the  clot    that 

:  'fccli'd. 

■'.  .1  'If licii  urn  ill  iirothniiiihiii.  in  the  iMindilion  known  as  •■melena 
■ 'latorilin,"  undouhted  licnelit  is  dcrivid  from  intra\cnons  in.jictions 
^■"m1  serum  or  hy  direct  Mood  transfusions,  prohahly  hccaiise  throm 

or  prothromhin  is  thus  furnished. 
■     .1  ih  fu-'u  nr-'l  of  fhr'-nil>"l'!'t^l'>'.     Since  this  suhstance  is  ijerived  fiiiiil 
ih  hloo.l  eells  and  tissue  cells,  it  does  not  seem  likely  th.it  a  .Iclic-ieiicy 


J 


If. 


Till,  iii.odii  AM)  Tin;  i.YMni 


I 
¥ 


iMiiiM  cNcr  (icciif.    ('I'ltiiin  oliscrvcrs.  liowcvcr     Morawitz,  I'lii'  oxaniplo 
liiy  jjifiit  strcvs  on  lliis  jis  an  iiii|iin'laiit   lactor  in  li('iiitirrlia<ji<'  «lisoas»'s 

4.  All  <  xrisn  of  (iiillthnimhiii.  The  iiii(|iiiil)t('(l  increase  in  tliis  siilistanci 
tliaf  ean  lie  In'oimiit  aluuit  exi)eriiiientally  liy  in.jeetin<;  liiriKliii  or  peji 
lone  into  aniiiials,  lias  stitiiiilated  earel'iil  sean-li  for  a  similar  increase  in 
the  l)ioo(l  in  clinical  eoiulitions  in  which  aluioiiiial  lilood  c|ottin<r  is  (in< 
ot"  the  syiiiptoiiis  (  \Vliij>plc"  1.  Aiitithi'oiiil:iii  is  sai<l  to  lie  increased  in 
septicemia,  pnciiiiionia,  miliary  tiiherciilosis,  etc. 

fi.  .1  (]i  I'll  i<  iini  iif  call  iiiiii  itiiis.  Altlioiitrh  at  one  time  it  was  siiiiposcil 
that  this  mi<rht  he  resi>oMsihle  Tor  tin'  feeble  clottinj;  in  liemo|)liilia,  ii 
has  not  lieeii  found,  after  \eiy  extensive  trials,  that  the  exhihitioii  I'l 
("a  salts  in  any  way  relieves  the  condition.  It  is  said,  however,  that  tli' 
slow  co;ii.'iilation  seen  in  ohstnietivc  .jaundice  is  decidedly  shortened  ]i\ 
treatment   with  calcium  salts. 

One  thini;  stands  out  i>rominently  in  connection  w  ith  tl;i'  whole  prohleni, 
and  thjit  is  the  close  relationship  of  the  Mood  platelets  to  the  elottiii;' 
])rncess.  l'"iom  these  cells  are  derixcd,  according  to  Howell,  not  only  tli' 
prothromhiii  luit  ;dso.  as  from  other  cells,  thromlioi)liistin.  If  is  not  su; 
prisinu'  theiefore  to  find  that  decided  alterations  in  the  i>latelet  com  ' 
occur  in  cases  of  faulty  lilood  elottiii};,  and  that  local  aceiiiiiulations  <•■ 
these  elements  within  the  Mood  vessels,  prodiK'cd  hy  their  cliimiiin;;  t" 
srether  or  aL'^rliitinatiiiir.  is  followed  by  a  formation  of  local  clots,  as  n 
throinliosis. 

Hemorrhagic  Diseases 

In   many  of  the  so-ejillcd   heinorrh.'ijiic   diseases   (acute   leuceniia   ani 
Jispastic   aiiemiji  I    and   in   the   heniorrha<,'ic    varieties   of  diphtheria   ar  i 
smallpox,  the  jilatclet   count   drops  from  its  normal  of  lietween  2(Ml.(Hii 
and  ,s(l(l,(((lO  per  ciiliic  inillinieter  to  well   lielow   10(>,()(H),  and  indeed   i^ 
these  conditions  it  is  fre(|nentiy  diflicuJ!  to  find  ;iiiy  jilatclets.    Siiiiiples  n 
Mood  clot  outside  the  liody  within  the  normal  time,  hut  the  clot  is  sn: 
and  usiiiill>'  fails  to  retract   in  the  iioriiial  manner.     It  is  on  account 
this,  rjitlier  than  slow  el.ittini;  that  the  hemorrhas;e  contimies,  so  that 
aiipraisiny  the  gravity  (d'  the  symptom  it  is  liest  to  measure  not  the  cl' 
tills."-  time  hut  the  time  that   it   takes  for  lileedin;.'  to  cease   from  a  sin 
skin  wound,  as  in  the  lolie  of  the  ear.    This  can  he  very  accurately  d" 
liy  applying:  Motlini.'  paper  at  reirular  intervals  to  the  puncture  (I)iikc' 

The  most  iiitcrestiiii,'  and  jit  the  same  time  the  most  mysterious  of 
conditions  in  which  lilood  clottiii<.'  is  interfered  with  is  In  mafihiliii.  "\ 
elottinjr  time  is  loiii,'cr  than  normal,  hut  even  after  tlie  clot  forms,  hli' 
iiijr  is  likely  to  continue  hecaiise  the  clots  are  very  readily  displaecil.  i; 
clotfinjr  time  and  lih'cdinj.'  time  are  increased.     .So  far  no  chanire  in  ' 


lil.OdI)    (i.OTTIXO 


113 


(•lc(ftin<i  factors  of  llic  lilood  liiis  Itccii  (Icnioiisf rated  in  this  discasp;  the 
,  (iipusclcs  and  tin-  ])iatclcts  an-  normal  in  nunihcrs,  filinno(;cn  and  eal- 
.iiiiii  salts  aro  noinial,  and,  as  Howell  has  shown,  there  is  no  excess  of 
.■,iitithrond)in.  One  sifrnilieant  fael.  however,  is  that  the  addition  of 
ihronilioplastin  or  of  its  aetive  inirredient,  kephalin.  jrrpatly  shortens  the 
cl.itlin','  time  of  the  hlood  when  it  is  removed  hy  veniinineture.  Tn  ajrree- 
meiit  with  this  oliservation  it  has  lieen  found  that  heniophilie  hlood  I'lots 
iniich  more  rapidly,  indeed  sometimes  in  the  usual  time,  if  it  is  allowed  to 
(|(,w  over  eut  or  damau'ed  tissue  and  so  heconie  mixed  with  tlinnuhoplas- 
liii.  These  faets  taken  tojiether  would  seem  to  indieate  that  the  fault 
must  lie  in  a  delit-ieney  in  jjrothniniliin,  and  slnee  this  is  derived  mainly 
iVom  the  platelets,  whieh  however  are  not  d<"creased  in  niimlier.  we  must 
lurlher  assume  that  these  elements  have  nndert'one  some  (pialitative 
ilian.'e  preventinsi  their  disinteirratioii.  An  aeeoiiii)anyinir  defeet  in 
tlieir  ai."-rlutinatiiiii-  properties  would  at  the  same  tiiiu'  explain  their  fail- 
ure in  hemophilia  to  eluniji  toyelher  at  the  site  of  the  heniorrha-re  so  as 
In  liloek  till'  smaller  vessels  witii  thromlii:  heuee  the  proloUL'ed  lileedinu' 
time  even  after  eloltinu'  has  oeeu     ^m1. 


Thrombus  Formation 

'riie  liist  fornu'd  i)oitioii  of  a  thronilius  is  i>aler  than  those  formed  later, 
l.icanse  it  eontains  excessive  nundn'is  of  i)latelets;  and  it  seems  clear 
!li,il  it  is  hy  airulntination  of  these  into  masses,  which  then  slick  in  the 
i. 1(111,1  vessels  and  liv  disinte-rratiny:  slied  forth  i>rollirondiin  and  throm1)o- 
|.l,istiii,  that  the  clo11iM<:  starts.  This  plat. 'let  au'U'lntination  may  result 
niiiii  sta-iuation  in  the  hloodllow,  or  from  rouy:henint:  and  danni>.'e  to  the 
- . --ei  walls.  Stauimtion  may  he  due  I'itlier  to  failure  of  the  circulation 
,is  a  whole  as  in  heart  diseasi'.  oi-  to  local  ])h\sical  ;ilterations  in  the  vas- 

■  ul:ir  tuhe.  setting'  up  conditions  in   which   e<ldy  currents  with  stau'nani 

I Is  of  i>l 1  are  foriiie<l,  such  as  will  occ\ir  at   jilaces  where  the  vessels 

-  i.l.lcnly  li( MIC  wider,  as  in   varicose  veins,   in  aneurisms  and   at    tlie 

-  i.l.leii  hend  of  lar!,'e  veins.     The  first   forim'd  (  i)latelet )   thiomlius  is  fol- 
..■•r.l   hy   one  of  a   darker  color,   which   lills   tlu'  vessel    uj)   to   the   lu'xt 

iiistnmoiie  hrancli.     Similar  stajination    may   also   follow   the   ohstruc- 

■  .11  caused  hy  loil<rmclit   of  emholi    in   the   smalh-r  vessels   (air.    I'orciu'u 
'    -lii's  in  line  suspension,  hacteria,  etc.i.     The  thromlti  in  such  cases  are 

■V  small  and  occur  particularly  in  the  capillaries  of  the  liver,  spleen. 
I  lunu's.  The  small  throndii  often  serve  as  foci  from  which  eloltiii!-' 
vads  into  the  lamer  vessels,  this  heinir  ofttMi  encnuraired  hy  an  increase 

tin a-rulahility  of  the  hlood.     When  the  iutima  is  inllamed,  it  is  jios- 

-Mc  tliat  excessive  amounts  of  throtnhoplastin   are  pi'odu I  and   that 

'  i^  neutralizes  the  .-mtitlirondpin  in  hloocl  mo\  iii'.' so  slow  ly  tliat  it  is  not 


I 


in 


•iiii;  iii.(i<ii>  A\i>  Tin;  i.VMi'ii 


I cpl.-ici'il  li.\  I'li'sli  liliiiicl  hfliiif  I'liiltiMi;  I'lisiii's,  HI'  it  liiiiv  Id'  tliiit  silli 
sliiiiccs  ilcii\<'i|  riiiin  till-  iiill;iiiii-il  tissue  cause  the  platelets  to  a(,'s;luti 
liate.  The  iiii'ieasetl  eiolliiij;  iilliMi  (ilisei\  I'd  after  tile  iiijeetinii  i»|"  lieliin 
lytic  airciicies  i  loieiyii  seia,  snake  .ciioiPi,  etc.)  inav  also  lie  (iiii'  Id 
platelet  a^u'lllt illation.  Like  tlie  tliioiiiliosis  t'ollowiiiij:  eiiiliolisiii,  tlic 
I'ldttiii'^  oeeiiis  at  lii-st  in  tlie  capillaries,  tlie  initial  tliromM  containing; 
masses  of  platelets  aloiiy;  with  skeletons  of  lilcoil  corpuscles  and  celK 
froMi  tlie  lilooil  fill  mini;  oiiralis. 


ClIAITKi;   XIV 


I.VMI'II   Fdl.'MATIdN   AND  CIWCn.ATH  )N 


i< 


ta 


GENERAL  CONSIDERATIONS 

LynipliMtics  iili'  iiiciililii'<|  veins.     Tlicy  jjriiw    rioni  the  veins  in  iMiitny- 
oiiie  life  iis  lintis  of  I'n.liiiiieiimii.   wliieli   iife  lirsl    visilile  in   tiie  liiiiiian 

eiiilil-Mi    in    tin-   sixth    week    uf   (1<>\  elci|iirient.       'Die    eiilliest    onl  UTou  til    ne- 

cui's  from  tlie  inteiiinl  .JMV'ubir  vein,  :inil  tlw  enilotlieli.il  liuds  somi  lie- 
eiiiiie  Imllow  Jiinl  .join  t(i<ictliei',  l'i)iniinir  tirst  :\  ])le\iis  iinil  suliseqiientlv 
M  s;ie,  li'oni  wliieli  ;iLriiiii  lyin|>li;iti<'  vessels  niiide  df  enil"lliiliuni  u'nivv 
.lilt  to  inviiile  tlie  skin  of  iln'  lieml.  neek,  tiioiiis  Mini  mill,  iiml  ]>iirtly 
the  deep  stl'lletllfes  oi'  tile  iie.id,      'I'iie  siie  is  lilt  ililMteiy   t  |-.llist'oillied   into 

LMMiips  of  lynipli   uliinds.      At    a    Inter  sl;ii:e  simil.-ir   les  develop    from 

eeit;iiii  of  tlie  .ilpdoniinnl  veins,  foriiiiim  :i  ret  roperitoiienl  s;ie,  from  vvliieii 
'.;rov\  out  llie  lympiwities  of  tlie  iilidomiiuil  ;iiid,  to  :i  i-ertiiin  extent,  of 
the  llior.Mcie  viseeiii.  A  similar  pair  of  saes  also  develops  from  the  iliiic 
veins  siipplyinu  the  lympliaties  for  the  skin  of  the  le'js  and  alidoniiiial 
walls.  The  retroperitoneal  and  iliae  saes  then  heeoiiie  eoiiiieeted  with 
the  .jiil-'iilar  sac  liy  means  of  the  thoiaeie  duel.  In  the  eiiihrvci  there  are 
no  valves  in  the  lymiihatie  vessels,  so  that  the  whole  system  can  he  in- 
.jerted  cither  from  the  thoiaeie  duel  or  from  the  skin,  sliovviiiir  (dearly 
tliat  the  siiperlieial  and  deep  Iv  iiiiihatics  are  parts  of  one  closed  system 
nf  vessels. 

Anatomists  have  suci'eeded  in  iraeinu:  the  course  of  the  lymphatics  in 
iriaiiy  parts  of  the  liod.v.  This  know  ledue  is  of  irreal  impintaiiee  in 
liiniieclion  with  the  spread  of  infections,  etc.  I.vinphalies  arc  ahini- 
•hiiit  in  tlic  skin,  the  intestine,  and  connective  tissues,  lint  arc  aliseiit 
iioMi  ih(.  niusi'le  hiindlcs.  from  the  hepatic  lohiiles  ilhoiiu'li  present  in 
ill"  eoiniei'tivc  tissue  hctwceii  theiir,  from  the  siilistjiiee  of  the  spleen. 
:iMd    from   the  central   nervous  system. 

Tlii    lympliaties  liavi'  tin-  same   functions  as  hlooil   ea|iillaries,    namel.v, 
:il'soili  siilistanccs  fi'om   the  tissue  spaces.     There  is  sonie  evidence  to 

'  "*\    that   this  ahsorptioii  may    he  selective.      When   in.ji'ctioiis  are  made 

•''1  the  iH'ritoneji!   I'avjtv.  llu'   pathwav    of  alisoi  lit  ion    niav   he  either  the 

" 'd   vessels   or  flic   lymphatics,    accordinir   to    the    nature   of   the   stih- 

t.iliee  injected.     Tiuc  solutions  arc  .ihsiuhed  hv    the  lilood.  hut   maiiulcs 


J 


i  J.'  ^-•'' 


lu; 


Tin;  isi.iiiii)  AND  •nii;  t.vmcii 


iiic  tiikcii  up  li>  sjx'ciiil  liULT*'  <clls  slidw  ins;  |>liii«;iir\  tir  poui'is,  anil  trariv 
I'lTicd  to  till'  lyiiiplijilics  I'lif  cxariiplc,  tlinsc  nf  tlic  diapliiiiLrni.  A  siiii 
ilar  sclfclivc  alistnplidH  is  well  known  in  tlii'  case  of  tiic  villi  «(  ilic  in 
tcstint',  wlii'ii-  I'at  i)asscs  intn  tlic  laclcais  ami  carlMiliydralcs  into  flu 
hliMid.      It    ai)p<'ars  as   il'   lyiiipliatic   adsoipli<ni.   Iiotli    of  solid    iiialt'iiaN 

and  of  sohitions,  iciprii'i's  tlic  ( jn'iation  of  jilianocx  tic  ci'lls. 

The  ih'wcr  conception  of  tlic  1\  nipliatics  as  a  closed  svstcni  is  at  \aii 
ancc  with  tlic  older  one.  in  wliieli  tliix    \\iTe  supposed  to  irel  snnillcr  aii'' 
smaller,    and    tlicir    walls    less    and    less    complete    until    ultimately    the 
fadrcl  (itV  into  ihe  tissue  r.paces.     These,  how cx cr,  hear  mi  closer  I'clatioh 
ship  to  lymphatics  than  they  do  to  lilood  capillai'ics.     The  tissue  spai-i 
incluile   all    the    minute   spaces   lielwecn    the    tillers   and    cells   of   the   cie 
rii'iMixc  tissues  anci  lietween  the  pareneli.\  in:i  ot   Ihe  cnuans  am!  the  '^n-.r. 
serous   cavities    of   the    liody    (pleural,    pcrlti  ncal  ■,    as    well    as    spceiall; 
dc\clope(l    tissue  spaces,   t'ormiiiir  the   suliarachiioiil   spaces  of  the  Iiraii 
the  sc;il;i  vcstilmli  anil  lympani  of  the  cichlea  ami  the  anlciiov  ehaiiilM 
iif  the  eye.      The   tlilids  in   lliese  spaces      the  tissue   fluids      arc  (luite  di; 
fereiit    from    the   lymph    in    t.lic    lymphatii's    lioth    in    composition    and    i' 
fuiietion.       Iinleed,    the    tissue    lliiids    arc    ainouir    the    iinist    varied    of   a' 
the    lluids    of   the    liod.v  .      The    spaces    iiniv     thcmsehes    liecnme    linUed    t. 
■.'ctlier  so  as  to  fi  rm  .-i  eirculatoi-.\'  system,  uliicli   is  (|nite  indepeiid'lit   i^ 
the  lymi'halics.     This  i-  particularly  the  case  in   the  lir.-iiii.  w  hcie  the  tl- 
sne  spaces  surroiindin','  e\ery   individual   nerxc  I'cll  e\tend  into  the  siil 
arachiioid  ai'ea.  \\liere  they  drain   into  the  cerehral   siniisis  thiouyli   tl' 
araclmoidal    \  illi,    \vliicli    exist    as   lace-like   piojeetions   of  the   arachimi, 
into  the  dural  sinuses,  lieinix  covered  ]>y  a   layer  of  nn-sothelial  cells  sp. 
cially  aliuiidant   at   the  tips  of  the  villi,  where  they  foiin  cell  imsts.     (i 
scr\a1ions  of  the   passaye  of  sulist.ane  s    in    solution    liy   llicic    patliwa; 
h.ive  lieeii  made  liv  in.jcctiim'  i>otassiuin   fcrrocyjuiide  and  citrate  of  ii' 
into    the    siiliarachnoid    and    sulMliiial    spaces    and    afterwards    dctectji 
the  iircseiiec  of  the  salts  liy   mountiny:  seetions   in   acid    media,  so  as   ■ 
permit  jiriissian  Mue  to  develop,     (hdinarilv    the  precipitate  is  found 
or  near  the  villi,  hut  after  cerehral  anemia   it    I'oriiis  in  the  tissue  sp;i. 
that   surround  the  nerve  cells. 

There  ;ire  therefore  three  lluids  coliceinrd  in  the  transfcrem'c  of  I'l' 
materials  and  u'ases  lietween  the  u'ast  roiiitestinal  apparatus  ;ind  lin 
and  the  tissue  cells  namely,  tlic  hlood  plasnni.  the  tissue  lluids,  .ind  i 
lymph.  'I'lie  //\s/(.  //((/(/,  heinir  in  contact  with  the  tissue  elements,  sei  \ 
as  their  imiiieilialc  nntiitive  lluid.  and  it  is  ihe  function  of  the  lilood  ;i 
l.vmph  to  maint.'iin  it  of  proper  composition.  Kvcivtliinir  must  lie  Ira 
ferred    lo   and    from    the   tissue   cells   throuirli    the   tissue    lluid,    niakinv 


I.VMI'II    KOIIMATKIN    ASM    lll«  11,  ATK  l\ 


117 


iliiii'lori'  in  iii.in\  \\ii\s  tlic  iiiusl  iiiipurtiiMt  tit'  the  lluiils  (if  llu'  IhxIv. 
I'l  ill.'  tissue  fells  tlieliiseh cs  tliere  is  iilso  the  lliiiil  in  which  the  vjhIdus 
rMlliiiils  iiiiil  eiystiilloiils  thiit    enter  into  the  eiiin|iiisiti<iii   iit'  |ii'i'tii|ilasni 


ilissiiK cil.     This  e;m  he  i('iiii>\ec|  lioiii  eells  (iiilx 


•hilM 


leal  irieans, 


>iii'li  as  trriiiiliii!;  with   tine  saml   in   a    nnntai'  ami   snli,ieetin<;  the   mass 
III  a  pressure  iit'  several  Ihnusanil  atiiii>,s|iheres  in  a  livtlraulie  iliuehnei'i 


I' 


I'ss.     This   is   l<ii(i\\n   as   the    I 


IS.1III      JIIK'I  . 


The    ultimate    exehantre   ol" 


l' Ishill's  (leeiiis   lietwecii    tiie   tissue    lluiils  and    the   tissiu"   jiliees   aentss 

ihc  ecll  Miemliraue.  The  estent  ami  charaeter  nt'  this  e\elian'je  (lepen<I 
iiH  iiian,\  eiieumstanees,  some  alVeetiny;  the  eell  wall,  iithers.  the  (isiiiittie 
iikI  other  pi-dpelties  ot'  the  two  lluiils.  ( Mi\  iiMlsiy,  the  flinetiiili  (if  tlie 
I'll  iMllalinii  is  tn  maintain  the  tissue  llu'<ls  ni'  enll-eet  eiimpiisiliiiri,  the 
I'lniid  |)lasma  serNiny  fit  eairy  I'immI  uialeiials  and  dissnixed  nwirtMi  tn 
ihriii  (see  pai;e  oSO).  hilt  heim^r  assisled  in  the  iip|piisite  ruilelinn  cd'  Te- 
iiiii\  al  111' elTete  plinluets  li\  the  l\  mph.  The  lymph  is  purely  a  sea\eni;er: 
till'  lijiiiiil  is  liiith   piir\e,Mi|-  and  sea\ ciiuer. 

The  aliiive  desil  iptiiiii  iif  the  ly  nipliat  ies  is  nut  linix  ersilly  aeeepted 
ii\  aiialiiliiists,  eei'tain  <i{  w  limn  lielieve  that  the  l\  mphat  ies  are  de\  ehipeil 
iMiiii  tissue  spaees  and  ale  eiiiisei|Uent  ly  iiilleh  iiiine  extensive  than  they 
appear  111  he  f'riim  injeeted  speeimeiis.  The  almve  ennelnsidiis  are  hased  nil 
I  eeiiiistriietiiiii  miidels.  made  I'liim  serial  seetimis  id"  emlir\ miie  tissues. 
Ml   w  hieli  the   lymphalies   Ireipieiitly  appear  as   isnlated   vesieles   withoiit 

M^ili! iineetiiiiis.     The   failure  nt'  injeetiiins  tn   penetrate  iiitn  the   re 

iii'iler  parts  nt'  siieh  a  lymphatie  svsti'iii  in  the  emliryn  is  altrilnited  tn 
I  111'  disciiMl  inuity  ot'  spaces,  which  is.  hnwcxcr.  remnxed  at  later  staires 
'if  dcvehipmciit. 

The   liiaiiner  id"  alisnrptinu   nl'   injected    llilids   dues    lint,   hnwcver,   slip- 

I'l'i'I    the  view  that    the  lymphatics  are  directly   eniiiiectiil    with    the  1is- 

^iii'  spaces.      When   all    the   striietures  nt'  a    |iart    are    liiiatcd   except    the 

111,'iiii    artery   or   vein,    in.jeeted    pnisniis    which    alTeet    central    structures. 

■I'-li  as  the  iierxe  centers.  develn|)  their  actii.iii  as  ipiickly  as  in    the  in- 

'  -'I  animal   (e.ir,.  stryehniiie ).     Similarly,  when   iiiirmelits  such   as  iiieth- 

'  ''  111'  liliie  are  injected   iiitn  the  ]ileural  cavity   or  sulieutainnusly,  they 

•piar  in  the  urine  hiiiir  hel'iire  the  lymph   nf  the  tlinracie  duct.      Such 

^mIIs  indicate  the  ])athway  nt'  alisnrption   to  lie  the  lilnnil    rather  than 

■'     I.Miipli    vessels.      Thmuirh    this    liitter    clianm>l    alisnrptinu    inneeeds 

'c  slowly,  hut  can  lie  iireatl.v  assisted  liy  iiiassairini:  the  site  nt'  iiijec- 

II.     When  colnred  snliitions.  sindi  as  India  ink  or  earmine.  are  injected 

'■"•ilt.Mlieously,    liiiwever,    a     very     JiiTl'ect     injection     ni'    the    nejirhliorill',' 

I  Iiliatics  ma.\-  ultimately  iieeur,  and    tlirnll';li    the  same    palhwa.vs   mi- 

'"iLMiiisnis  spread  fnuii  an  iiit'ected  area. 


i 


lis 


nil:  111  iMiD   \M>  Tin;  i.ymi'H 


EXPERIMENTAL  INVESTIGATIONS 

It  li;is  pidvtil  II  must  (liriiiiilt  priiltli'iii  tn  u'iiiii  ;iiiy  cxiicl  kiniw  Icdi;. 
cif  iIh'  iiriMliiriiiiii  111'  I  \  II 1 1  ill  liy  i'\|MTiiiifiil,il  iiH'iiiiN.  St.'iiliiii;.  sniiic  yi'iii  ^ 
(i^ii.  ill  icpi'.iliiii.'  iiiiiiiy  111'  till'  r\|)i  riiiii'iils  ut'  hIiIit  |)liysiiiliii;ists  ill  tin 
li'-'lit  111'  till'  liiwiT  r.ii'ts  III'  |i|iysir,il  clli'lllisliy,  iii|i|i'i|  iiiui'll  tllilt  in  ul' 
inti'ii'st,  aiiil  it  is  cliirilN  «iili  liis  wmk  tiiiit  \\c  will  ciiiii't'rii  oiirst'lvcs 
here. 

'I'lic   lllii'i|ll.'il    l>iM|>li    sii|i|i|y   of  ililTi'iiiit    ii"_niiiis   111'   tlic   liiiily    is   stiik 

iiiirly  ilii isliiiti'il   li\    riiiii|);ii'iii'_'  llir    l.\iii|ili    (luw    rrmii   Itii-  lynipliiitir^ 

lit'  tlir  liLT  \\itli  tllilt    t'ruiii  till-  tliiir;ii-ic  iliict.     No  lyiiipli   llii«s  from  tin 

I'lirillfr   llllli'ss   tile    iiillsi'lrs   ;i|-i'   tliriiwil    ililn   ;irti\ity    or   till-  liliinil    is    pIT 

vciitril  t'riiiii  liMNitiir  till'  liiiili  liy  li!.Mtiirin'_'  .ill  tlii>  vi'iii'<.  ( 'liiiiiL'i's  in  iIm' 
Mi-trriiil  liliiiiil  pii'ssiin-  ilo  nut  nlVi'it  tl:i'  tluw.  On  tlic  iitliiT  liiinil.  ii  trrr.il 
ini'ii'M--.'  in  tlir  ijim  I'lniii  tlii'  tlnn-iii-ii'  iliin  i-;ili  li'.'iilily  In'  iliihliTil  li\ 
ilistiirlijiiiri's  ill  till'  liluiiil  supply.  (  ilistnii-tiiiii  ul'  tin-  |>iirt;il  xciii.  fn; 
lAiiiiipli',  iiiiiiirili.ili'lv  iniTi'Msi-s  Ihr  lymph  Mow  fniir-  111'  tivi'  timrs  lici-jinsc  ul' 
M'lliills    i-iiiiL'i'slinii     111     ||ii.     intrsliii.-il     f;i  pi  1 1;|  rics,     wliilst     ;i     still     irii'iitr' 

iiM'iT.isf  piili.-ips  ti'iit'iilij  is  iiiiliii'i'il  liy  iilistiiii-tinii  to  tlic  iiil'ciior  vrn.i 
I'.iv.i.  wliii-li   liiisi's  till'  I'.-ipilliiry  pirssiiii'  in  Imtli  the  li\iM-  iiiiil  tin-  intiv 

tiliis.      Al'liT  liiiMli r  till'  liipjitii'   lyiiil>liiilii's  (;it   tlic  licpiitic   licilii-lr 

iikstinctiiiii  111'  III.'  \cii,i  iM\ii  nil  liiii'.'cr  c.iiiscs  the  oiitllou  of  l.\  iiipli  t" 
i.icrc-isc,  iiiiliiMt  in;;  llnil  the  l.\  iiipli  in  the  l;ist  iiicntinncil  cxpcrimciii 
iiiiisl    hinc  Clinic  rnnii  the  hepiitic  lyiiiph.-ilics. 

'riiese  iisiills.  Ml  r,ir  ,is  tlie>  l'h.  ciililij  lie  s;it  isriietoiily  CNpliiiiiCil  nii 
the  li;isis  tllilt  lymph  runinilinn  is  ;i  lillrdliuii  piocess,  thiit  is.  ;i  pmci  s- 
licliciiiliiit  llpiili  ililVelcncc  ill  iiieeli;inic;il  pressure  licluecii  the  liln.MJ 
e;ipill,irics  :ilii|  the  tissue  spjirrs.  The  ly  iiiphlll  ics  wmilil  then  sei\|.  :i- 
elninnels  ii,  nliiin  this  lliiiil  In  ih,.  Iihinil  vessels  lliriiii;;li  the  tliuiiici  ■ 
iliK-t.  'I'lie  ililTeiTiii-i'  ill  the  ni;ii:ii it ikIc  lit'  the  iiiei'cMsi'il  Ivinpli  llnu  I'riii 
illcle;ise    ill    iMpiihiry    piissinr    jn    ililTci-elit     regions    Wiinlil    lie    ilcpemlii  ■ 

nil  the  peiiiiciliility  1 1 1'  the  JiltiT,  tl ;i pi  1  l;i rics  ul'  the  linilis  liciiiiz  iiiiii  ' 

less  pci-iiicalih'  tlnin  Ihiise  nt'  the  intistine.  iiiiil   p.-iiM  icnlnrlv    uf  the  li\( 
Ainther   I'mcI    ill   ciiiil'm  iiiity    with    this   \  icw    einiccrns   the   eompiisit  inn   ■ 

the     l\niph     llnlii     till.    t\Mi     ir'jiiills,     th.-lt      I'lnlll     the     lilllll     l,\  mpll.-ltics    iieii: 

1' ■   in    pinliiii,    v\heii-;is   llnil    I'r the   t  ImrMcic   ilnct    ijoes   nut    I'iill    I.. 

Ill  liiml   llic   111 1    pl;isiii,i   in    this   rciijiril, 

.Mlhnnirh    lilt  i;i1  inn    in.i.x    expl.iin    the    ciinsiilci-iililc    iiiercjisc    in    l.vini' 
|!iu\    prnilM<-'<|   l>y   (  vtii'tlic  "l-,ilii:-'s   in   e:t|.i!hir\    )>t'essutc.   it    \n    im   iin:: 
siifliecs   In   cxpliiin    lymph    loiimil  inn     iimler     less    jilmofniiil     coiiilitin!  - 
When  .-i   muscle  m-  -.y   yhiiul   is  ;it   rest,  it    pinijiiees  priicticallv   111!  l.viiip 


I.VMI'II    FOIIMATION    AMI   <  ll((  I  I.ATIIIV 


11!» 


lull  (liiriiif;  iicfivity  tin-  llow  I (uncs  iiiiirkcil.    Tliis  cini  imt  lie  r\|>l,iiiit'(l 

h\  liltriitioii,  but  iiiiiv  !)»■  iicciiiintcil  for  by  ii  pliysicoclicniifal  |iiiiccss  - 
ii.iliirly,  (isiiiiisis.  Tin'  ciH'i'iry  i'ri|uiri'(l  l"(.r  tlic  iiftivity  of  tlic  lissiii' 
cell  is  proillK'i'il  l)y  clicniirjil  I'llilll^rt'S,  wiirrcliy  1iiI'J.M'  lllnlccllli's  liccniiif 
lirokni  down  into  iiiiincrous  siiiallcr  ones.  Tlicso  siiialU'r  iiiolccnlcs  iirc 
Ihcii  (lisclmrtri'il  into  tlio  surroiindin!;  tissue  lluiils,  tiic  osmotic  iircssurc 
of  wliii'h  they  increase,  uitii  the  eonse((uenee  that  water  is  atti'actrd 
li,\  i>sini>s.is  from  tlie  phisina  in  tile  lihiod  ca|iillaries  !see  \K\<ir  4).  This 
increases  the  voluiiio  of  tissue  llnid.  whiidi  is  then  drained  away  hy  the 
iynipliaties.  The  increase  in  nn)lar  eoneentration  will  also  alTect  the 
tissue  juici's,  tendinj;  to  nnike  the  cell  swell  uji  liy  alisoiliinji  water. 
In  lilaiul  ceils  this  extra  water  is  immediately  extruded  to  form  the 
wali'r  of  the  secretion  (see  pau'e  4^1). 

An  analo<;o)is  method  of  lymph  formation  is  not  conlined  to  sitimtions 
where  the  capillaries  ai'e  relatively  impermealile.  for  it  also  occurs  in 
the  liver,  the  lymph  (low  from  which  is  >,'rcatly  increased  l>y  the  injcc- 
lioii  of  hile  salts.  A  similar  process  no  doulit  residts  from  muscular 
activity,  altlKiu^h  in  this  case  the  tissue  spaces  must  form  a  continuous 
system  of  their  own,  Ihi-re  liein^r,  accordinir  to  most  authorities,  ntt 
l.\  niphatics. 

I'onsideralile  interest  has  liecn  taken  in  the  stimulating;  etVcct  which 
certain  chemical  suhstaiu-es  havi-  on  the  seci'etion  of  lymph  from  the 
thoracic  duct.  These  so-called  IjiiiiiiliinitxiiK  a  lieloni;  to  two  classes 
ciystalline  and  colloidal.  Of  the  former,  !,'lm'ose,  urea,  aiul  sodium 
cliloride  in  hypertonic  solution,  are  the  hest  known.  Stai'lini,'  explains 
their  action  as  dependent  upon  an  increase  in  the  osmotic  |)ressure  of 
ihc  Mood.  This  attracts  water  into  the  Mood  from  the  tissue  juices. 
:inil  leads  to  an  hydremic  pletlnua,  with  a  consei|iu'nt  ini'rease  in  eapil- 
Ijiy  pressiii'c.  If  the  lilood  pressure  is  lowcicd  liy  hemorrhajic  liel'oi-i' 
\\<r  hypertonic  solution  is  injected,  very  little  stimulation  of  lymph  Ih'w 
'"i-urs,  hceause  there  is  no  availaMe  lluid  in  tin'  tissue  to  protluee  the 
I'lrtliora.  This  oliservation  does  not.  houexci-.  vcr\  strouLrix-  siippoi-t 
til"  c\|ilanation,  lieeause  S4i  many  other-  distrrrliarrccs  ma\  resirit  frorrr 
i '  iiuii'i'ha;;e. 

The  colloidal   lympliajroirnes  iiiclirde   w;itcry  extracts  of  the  dried   tis- 

r.is  of  leeches,   crayfish,   and   mussels,   as   well    as   comiuer-i-ial    pe))torre 

'I'i  i\    proltalily   act   liy  darrrairiiifx  the   eridotheliirnr   of  the   capillaries,   so 

!l..it   liltratiorr  occ\irs  mor'i-  readily.     Althouirh   their-  miction   is  .lisplayed 

■  ;■(•  itiirticularly  on  the  !\  tuplialics  of  the  liver  and  int.-siines,  it  is  .-dso 
iipii^ii-erit  <in  the  skin  capillaries,  producing'  eutarreorrs  edeiira  ;ind  the 
I  . •■Illation  of  Misters  (nettle  rash). 


IL'O 


Tin;  iii,(Mn>  AM>  Tin:  i.vmiii 


EDEMA 

With  NlH'li  ill!  ilii|ii'i  iVi't  Iviiipvv  Iciluc  i'ip|nTiiiiii!»  tlif  |iliVNiii|i));y  "I 
lxiii|ill  ri)riiiiilii>ii,  il  is  iinl  Nlilplisilii,'  tliiit  the  ciiusfs  iil"  f\ffs.sivc  lli'fii 
iiiiil.iliiiii  1.1'  lliiiil  ill  iiiiil  littwccii  llic  lissii.'  iliinciils  nIi<>iiI<|  lie  little  iiii 

il<'isl I.     All  lit'  till-  (•iiiiililiiiiis  wliii'li  iiiiM'  lii'cii  iiinitiiiiii'il  iis  i-jipiilili 

111"  i-iiii>iMir  ,111  iiincii.M'iI  si'iTrlimi  i,(  l\iii|i|i  mu'Ii  us  iiificiisf  in  ciipilliiiv 
pi-i'sslll'i',  ll.\  ijli'liiir  pli'tliiilil.  .-irtiiili  111'  pdiMiiis  nil  the  «'lli|iilllt'lilllll  ai< 
likflv  III  ciiiisc  ciifiii.i  \(  the  Iviiiplialii-s  iit'  llir  purl  jirc  siimillimrnusli 
iilistlllrlril.  'I'll  pniiillci'  in  Illlilll.'lls  nlrliia  111'  tlic  Miliciilaiicdiis  tisslli'^ 
liUf  that  nlisi'iscii  rliiiicall\-.  it  is,  Iiuwi'MM',  ncci'ssaiv  lliat  llic  vasi-iihn 
liisliiiliaiicc  lie  acciinipaiiirii  cilhiT  liy  hiral  (lama'.'i'  to  the  capillai  \ 
I'liijiithcliiiiii,  siicji  as  is  pi'iiiliici'ij  liy  arscnii'  <ir  iiraniiiin:  nr  liy  a  iri'ii 
I'lal  tnNcniic  ciiiitlitiiiii,  such  as  is  si-f  up  liy  iifpiiiilis.  Whrii  laitn 
aiiKiiliits    lit'    saline    siiliiliiin    are    iii.jeeleil    iiitrax cnnusly,    exteiisi\c    e\ 

tia\asatiiiii  nf  lliiiij  iiia\    iif  iiitn  the  liver,  peiitmieniii  ami   iiilestiiial 

liiiiieii.  withiiiit  any  siilieiitanenns  eiji  ina. 
('iinieal  edeiiias  are  nf  at    least   tliii'e  t.\pes: 

1.  The  iiillaiiiiiialiiry  eileiiias,  in  whieh  the  llniil  permeates  the  eells  ni 
tlie  itilhiliieil  area  ami  dues  nut  shil't  to  other  Jiarts  nt'  the  liiniy  nilih  ' 
the   iiilliieMee  nt'  trravilx. 

2.  The  iiephriiie  eilemas.  in  whieli  tlie  lliiiii  is  nmre  nr  less  Innse  in  tih 
siilienlaneuns    tissues   ami    reailily    eliaiiires    its    pnsitinn.    ami    which    i. 

aeeiiliipailieij  liy  excess  iil'  Water  in  the  hloiiil  with  a  cnrrespninliiiif  ill 
crease  nf  siiiHinii  chlnriije;  tlie  )iercent;i};e  enncentratiiin  nf  sniliuni  chj.. 
>■'''''  ill  'Ik'  I'I'ii"!  remains  nnelianu'cil,  Inil  that  the  nthcr  coiistitiieii' 
iliminishcil. 

'■>.   (■anliac  edemas,  which  are  alsn  hypiMalie.  Init   are  nnni mpaniiii 

liy  tdnniu'cs  in  the  rel;iti\e  aniiiiiMt  nf  water  and  sndiimi  ehhnide  in  tl'- 
lilnnd. 

The  secniid  and  third  \arieties  nf  edema  nia\  nf  enurse  lie  mure  ■> 
less  present  tnt;etlier,  for  the  kidneys  are  likely  In  liecnme  see.iiidarii 
all'ectcd  diiriiiir  \enniis  stasis. 

The  salt  retciitinii  in  nephritic  edema  is  \eiy  siLMiilieaiit.  As  e^ 
plained  elsewhere,  it  is  revealed  liy  enmparin'j-  the  daily  oulimt  nf  - 
diniii  chlniide  liy  the  urine  with  the  cnncent ratinn  nf  this  salt  in  I! 
lilnnd.  Less  salt  is  eliminated  than  would  lie  the  case  in  a  inirinal  ii 
dividual  with  the  same  iiercenta'.'c  nf  salt  in  the  lilond.  In  many  ci^i 
also  edema  can  lie  diminished  liy  wiihlinldinj:  salt  frnrii  the  food.  W'iil 
and  .)a\al  have  coiieliisivel.\  shown  tlie  relationship  of  retention  nf  w:ir 
ill  the  lindy,  as  .judired  liy  variations  in  Iiody  weij;lit.  to  the  hvilrem 
eoiiditinii.   as  judjicd    liy    the    refractive    index    of  tlie    lilood   serum,   an 


I.VMI'II    riiKMATIiiN    AM)   ilUrl  rATtiiN 


1_M 


tlx 


limit   III'  Mill    ill   llif  ilirl.      A    M'i\    I'liiisiili'i'tiltlf   iclfiilinii   of 


wiilfr  ii.siiiilly  (HMMiis  lici'dfc  llu'if  is  iiiiv  fviilrlH'i'  III'  rilriiiil ;  iinlrnl,  a^ 
,1  fcMiit  til'  uiviiiif  suit,  tlic  liiiilx  uciiilit  iiijiv  iiH'ifiiM-  t'riiiii  live  Iti  m-vcii 
mIii(;|'iiiiis  (10  lo  I'l  |iiiuiiiis)  uitliin  a  iia\  nr  Iwu  willmiit  the  appear- 
, I  lice  III"  piifliiifSM. 

Till'  calls)-  iif  tlic  cilriiui  iliii'in<;  salt    rili'iitiiui   is  iiu  ilmilit   i-lnst'ly   rr 
liilril  III  till'  lu'liiin  III'  l,\  inpliiiiiiiirui's.     In  a  imrinal  pcrsnii  cxfi'ssivi'  in 
■.Tsliiiii  III'  sail  is  iniiiM'iJiatrix    rnlliiucil  liv  cxrirliiiii  nl'  the  cnitss  tlii'iiilt.'li 
tilt'  kiilncv.      Wlii'ir  llic  Uiiliicvs  air  (lisciiscij,   tliis  cnci'ss  of  salt    is   re 
iiiiiH'il   ill   till'   liliiiiil.   raising   its  nsinntii-   jtri'ssiirr  anil   attrai-tiiiK   \Nat('r 
I'liilii  till'  lissllr  lllliils.     This  Irails  In  cxccssix  r  lliiisl,  tin-  illlliilicil   walcl' 
lifiiii;  liscil  III  rrplacc  that   Inst    I'rnni  the  tissues,      ilut  all  the  ci'n  statline 
l,\  iiipha;;iij.'lles  ilii  nut,  «hen   present    in  cNerss  in   the  lilmul  nf  nephi'itie 
patients,  neeessarily  eause  etleiiia ;   urea,   I'm'  example,   may   aeeiiniiilaie 
r.msiilei'alily  withmit   any  siieli  elTert.     The  (lilTeient   aelinn    is   usually 
.ittiihiileil  to  ineipiality  in  the  ililTiisiliiiity  of  the  Iwo  ei-,\  slallnids  thi'oi|i;|i 
minimal    iiiemliranes,   soiliiim    ehloi'iih-   ijill'iisin'.'    miieh    less    reailily    than 
urea. 

Il  is  most  inipoi'taiit  to  note  that  the  llniil  in  eijeina  is  loose  in  the 
tissues  and  eaii  he  ilraineil  ax\a.\  1p.\  the  inseilion  of  tulies.  There  is 
.'ihsoliilfly  no  eviilenee  in  sup|)oi't  ol'  the  elaiin  of  .Martin  {''iseher  that 
edema  is  due  to  imliiltition  of  water  liy  the  enlloids  of  the  tissues.  This 
'iHi'stiiiii   has  hcen   flllh'  disrUs.scil  rlsewhere   (pave  (i'J). 


I'.LdOl)  AM)  I-Y.Ml'll   UKKKHKNt'KS 

(Moiioirraphs) 

ll"».ll.  W.  II.:     Tlir  lliirvrv   l,...tmvs.  .1.   ji.   I.i|.|.i,ir,,|t   (  k.,   \ii.  L'7:.'. 
^l:>rliii-.   K.   II.:      lliini:iM    l'iiv.-<i(.l.i;;v.   l.m   \    Idiin.i,    I'.U.-,, 
I.'"",.,  \.  II.:     .VnJi.  int.  .\|,..|.,  I!il7.  \ix.  :.r,l 
'U  illi;Mn-iMii.  I',  .s.;      .\r,h.   Int.  .\l.il.,   llilD.  x\iii,  .'iH.'.. 
I'"»ir  :iiicl   lliTiii:      Tnif.  Sue.   liiol.  .-oi.l   .\li'.|.,    Cili'..   .wiii.  .'n.'i. 
i."""  :iiii|   l.'ulicrt.-idii:      .Iinir.   Kxp.   .\liil..    IHlii.   wiii,  Jl'.i,  L'.:'.i.   '111. 
n'.titlir,  <;.  (!.:     (^iiiirt.  .I.mr.  .M.'.l.,  IIHJ,  vi,   11.1. 

Ii"«rll,  \V.   II.:      if.   Il:irv,y    l,,iturf;   ;iN„   .\ii,.  .1 ■.    1'1,vm..I  .    liU:;.   xw.i.  •-''■■I. 

I'liiiKiT.  C.  K..  :iiii|   K.   i;.:  '  .\in.  .IiMir.  I'livsi.,1.,   l:iir..  xli,  .".. 

•  '■uh\    ;iri,l    .Miiiiit:       .\iili.    int.    .\|.-,1.,    llMi;,    \\  ii.    |li|;    .\ni.    .Imir.    l'li.v>ii.l..    UH." 

xxxviil,  l'.'!.;. 
\l'liN  T.:     (^iitirl.  .Ii.ni.  .M..!..  Illlii.  iv,  11. 

'  :iiiMciii  !in<l  .Mrmli'iilNill:     .\iii.  .Itmr.  riivsinl.,    li'll,  \xxi\.  L'-."i. 
H-"ill.  W.   II.:      .\r.lk    Int.   .Mnl.,   I'.HI.  xiii.  Ml. 

•  ' 'lii'.  T.  (i.:     .lonr.  I'liv^i.,1.    Is'ir,  xxi.   In:;. 

Vlii|.|.|,..  (i.   II.:      .\r.li.   int.   Mr.l.   I'.'lL',  ix,  :;i-..-i:  .linir.   Kx]..   Mi. I..   I'.HI.  xiii,   1:!!!. 

\  l.i|.|.li',  (!.  H.:     .\rili.  Int.  Micl.,  l!il.;,  xii,  li.'.T. 
:'iiiu.  W.  \V:     Anil.   Int.  Mnl.,   liili.',  ix,  L'."iS. 


J 


^jJt-Jf*^' 


I'AKT  Mi 
TIIK  CII^  ri..VII()\  Ol"  TIIK  HL(M)I) 

rii.\i>Ti:i{  xv 

Ill.odl)  I'li'KSSI  l.'K 

Tile  iilijcrl    lit'  till'   I'irrllljltinli    is   In   liiMilltMill    lIlliMI'.'li    tllf   lisslli'S   n   slip 

|>lv  I'l'  111 1   tli;it    is  .ii|<i|iiiit('   III   liiri'l    till  ir  ilcliiHllih    I'dl'   milriliiriit    Mil. I 

ii\>'_'<>li  .'iiicl  til  niiiiiM'  till'  tiTctt'  iHiHliu-ts  III'  llii'ir  liirtiili<ilisiii.  'I'lir  i\< 
liiiiiiils  \,iiy  ;irr(iiiiiiiv,'  t"  llii'  iirliv  ilii's  nf  tin-  tissih',  lii'iiii,'  pjii  lii'iiliirl'. 
\  ,iii;i)ili'  ill  till-  r.isi'  iiC  sllrli  tisMli";  lis  till"  inilscillill'  tiMil  llir  l.'liliiillli:ii 
III  sIiiiIn  iii'_'  till'  |ili\  NiiiliHjy  lit'  till'  riii'iiliitimi  \M'  liflM'  liirii'l"i)i  I'  ti>  lii';i! 

in    iniliij    two   ;l^|ll'l■t^   nl'    llii'    |)lii|ilriii ;    il)    tin'    fiillsi'    I'or    tin litiniliiM'- 

liiiiiiijliiiw  .  .'ilnl   I  2     till'   iiM'cli.'iliislii   li\    \\  liirli  iiltrr.it  inns  in   tliis  Ii|iihiI||iix\ 
;ilr   linillylit    :ilMiiit. 

It"   \M'   iiiiii,    Mil   ;il'li'ry    \\i-   sjuill    litnl    that    tin-    lilimil    i-si-iipi's    rinm    it 
I'liijif  sihIi    a    |iii'sviiii'   tli.it    il    is  tlii'iiwii   III  ,'1    In'iiilit   iit"  aliinil   si\    I'ri'l. 
tliiit   ils  until. .vv   i^  |.i  .i|iiii  limial  In  tin'  si/c  nl'  tin-  arti'iy,  ami  tliat  it  im' 
satis,      ir,   nil    till'   i.tlii'i-   liaml,    xw   i>im'ii    a    \('ili.    >m'   shall    (linl    tlial    tin 
Miiuil    «rlls   niit    williiiiil    any    \riy    r\ii|i'nf    pri'ssnri',   anil    tliat    it    llnw - 
in  .'I  I'liiil  iiHiiiiis  striaiii,  its  iintllii\\    lii-inii  Ilir  sanii-  in  a   nnit   nl'  linn'  ■<■ 
tliat   111'  llir  .ilti'iy,  |ir.i\  jiliil   tlir  two  vr.si'ls  al'r  tin'  nnly   inirs  supplx  ii   : 
till-    pailhiilac    ail. I        'I'lii'    unii'i-al    cnni'itiiins    i.'iivi'nijntr    tin'    lili«iimi'.\ 
ail'  till'  sanii'  as  tlm-i-  .j.imi  niir,'  tlir  llnu   nl'  llniil  tiiiiiii!.'li  any  s\striii  '' 
tiilii's.      |-'iii-  ixaiiipli',   in   till'  rity   watrr  mains  it    is  knnwn   to  I'vcry  n' 
that   thr  rati'  nl'  nntllnw  I'mm  aii>    p;irt  nl'  tlir  systi'iti  di-pi'inis  (inall.v  .>: 
I  v\  n   t'ai'tniN     il      till'  ililTri'i'ni'i'  in   picssni'f  at    llir  l>t'i;iniiin}r  anil  cinl  " 
thr  systi'iii,  aiiil   I  •_' i    till'  ralihrr  nl'  tin-  tiihi'  at    till'  niitli't.      W'r  ma.\    i' 
rlrasi'  tlii'  nntllnw    liy    lai^iii'.'  thr  pii'sslllr  at  tin'  liryilinini.'  <it'  thr  sys|i  i, 
till'    i-alilii'i'    nl'    till'    niillrl    riir.iiiw  hill'    irmaiiiinu:    rdiislant.    nr    \i  r    in, 
m.'iiiilaiii    llif    |irissiiii'   rniislani    hnl    im-rnasi'   the   r.'ililiri'   nl'    thr    nnH' 

III    Ihi'   I'iniihilinn    nl'   tin'    lilnml.    thi'    ililTi'li'liri-    III    pli'ssMI'r    at    I  hi-    ! 
i;inniii'/  ami  rinl  nl'  thr  riii'iilal  inn   is    rurnishi'il   hy   th.'  pnmpinjr  ai'li 
nt    l!n-   hiMii,  ;;ini    llii-  allnraliiiu    iij'   thn   ralii..'!'  iif  the   nlltlrt    i>   prnv;.!     • 
for  liy  till'  i-niistrii'tinn  or  ililatatinn  of  tin-  IiIoimI  m'sscIs.     Thrsc  <!itii|' 
physical  prini'ipli's  imliiati'  thi'  diicrtion  \Oiii'h  a  stiuly  of  tin'  rirculatn  m 

122 


if 


|>.|  IN)|I     I'lll  SSI    III 


V2\ 


-liipillil  tilk<v      'riiiN    ilHlif.ilr  IIimI    uiir  tiisl   I'dlisiililllliiill   sImhiM   lir  of  till 


iiii'JIll    iMuml    lilt'sMiri 


I    pit 


liiiw    it    i  tniiM'il,  ami   lii>\\    it   niii  lie  iiiailc  to 

\;\IV.       Alter    ur    Iimm'    Ii'.mmcii  «  c    liliiy    tliril    |il(ici'r<l    to    a    inol'' 

|i,iitiriiljii'  cvaininiilinh  i>(  the  iiHTliaiii«lii  •>(  tlic  |iiilii|i  lliat  is.  cif  tin- 
li.aitlM'al  ;  linn  (iiiallv  \M'  may  |ii(ircci|  to  f\aiiiim-  I'm-  liatiiic  iff  lli«' 
|iiiircssi>   li\     wliirli    till'   falilM'i'   III'   till'   aiti'rirs   is   rulltfnilril. 

THE  MEAN  ARTERIAL  BLOOD  PRESSURE 

Till'  lilst    Iili'l'nilliNilc   til  till'  ili\  I'stiltlltiiiil   III"  till'   liliiiiil    |il'''ssillr,   as  III' 
iiiiv    iillii'i'  pliysii-al   iniililt'iii,  is  that    \.  r  sliniilil   pnsscss  siimc   means  li\ 

viiiii'li  it  i-aii  111' i|iiaiilitatiM'ly  ineasiiiril.     Tlii- cat  lii'st  attempt  in  ai iii- 

plisli  tliis  was  maile  liy  flie  Kiiirlisli  seiiiitist,  the  \W\ .  Stephen   Hales,  a 

hltle  iiM'i'  a   eentiii-\    after  llarve>    pillilishnl   his  ai tint    nf  the  eireula- 

ti.iii  cif  llie  liliMiil  Hales  I'liniieeleil  a  L'lass  tiilie  nine  feet  in  IriiL'th  «ith 
,1  si'\ereil  artei'>  nl'  a  Imrse,  the  eiiniiei'tinii  lietween  the  t\Mi  liiiii!.'  inaile 
h\  means  nf  a  pieee  nf  lirass  pipe  jnilliil  In  the  W  illilpipe  nf  a  unnse  as  a 
-uli.liliite  fill-  riililier  tnliiii!.'.     lie  fiimid  mi  niitvinir  ilie  liyatnte  mi  the 

,iiii'r\    that   the  Mnnil   ruse  in  the  tnlie  In  a   heii-ht  nf  ei<rht    t'eet   ainl  till 

i'lihi's  alinve  the  h'Vel  nf  the  left  velitriele  nf  the  lieai't.  ami  that  when 
,il  full  heiyhl  it  rnse  anil  fell  \\illi  caeh  pilUe  Ihinii'.'h  a  ilistanee  nf  twn, 
llii'ii'  nr  fniir  iiiehes. 

Mercury  Manometer  Tracings 

The  sniiiew  IukI  elllile  lillt  very  siL'liitieant  experiimiil  nf  Hales  elearl.v 
•  ^talilisili'd  the  (Nistellef  nf  the  elinlMiimis  plessnit"  at  wllii'h  the  liiiinil  is 
inaile  tn  eiii-iilale  tlirnnuli  the  arteries.  Tn  rentier  pnssilile  a  further 
iii\  es|i<ralinn  nf  the  faetnls  mi  \\hieii  this  pressure  ilepends,  it  lieeaiiie 
liii'essary  tn  in\ent  snme  iiinre  ennvenieiit  means  I'm-  its  measmement, 
i'ilt  this  «as  lint  aeeniiiplished  until  a  eentlir.\  later,  «  hen  rniseuille  ap 
|iii(d  tlie  III)  riiirii  iiiiiiiitiin  li  r.  uhirh  l.ndwii;  siiliseipieiit  iy  adapted  mi 
ili.ii   iraeintrs  miirlil   lie  taken   i  l'"i'_'.  '_'li. 

Ilaviliji  liefnre  ns  siieli   a   traeiiii;  as  shn«  n   in    l"i'_'.   '-'"J,   let    ns  emisider 

!;"\v    it    liiav    lie  Used   in   the  sluil\    nf  lilnml   piissure.      The  first    lliini:  we 

'  i!|^l  dn  is  In  measure  the  a\eraue  lieiuhl    nl'  llie  liaeiiiv;   al>n\e  the   line  nf 

•in    pressure;    the    mean    arlerial    lilnnd    pli'sslire    is    then    eipial    In    this 

!i-t,i multiplied  liy  t\',ii,  heeaiise  the  dislanee  lhrnu<rh   whieh  the  mer 

'Miy  has  moved  up  in  the  liiiih  nf  the  maiinmeter  earryiiiLr  the  \vril- 
•  ".'  piiiiit  is  milv  line. half  nf  its  total  disjilaeemen!  Sinei'  niereury 
I  .iliiMit  l:!.ri  times  avier  than  an  e(|nal  vnlnnie  nf  Mnnd,  the  almve 
li.i  isureinent    must    he   multiplied   hy   this  lii;nre    if   we  desire   tn  e\|iress 


1 
J 

J 


^M^i^nt) 


124 


tin:  ciiiri  i.ation  ok  tiii:  iii,(»)i> 


out-  ri'slllt   ill  trims  .if  llic   lici'.'lit   In  \\llicli  till-  IiIiiihI   lili'NSllic  cnlllil   rai^' 

a  colniiin  nl'  IiIimkI. 

Ill   .•irtcrics  III'  .•i|)|Miixiiii;ili'ly   llic  saiiii'  .si/c,   (lie   mean   .-ii-tiiial    I'Io.mI 

lUfsMUT  (IcicN   not    iiiarkcilly   vary   in  dilTiTcni    m: iials.     Tims,   in   tin 

caiuliil  aili'iy  nl'  llic  diiy;  it  avi'iiii;cs  alioiit  I  III  In  I'JII  inin.  llj;,  in  tli;i' 
of  (lie  i-al  aliiiiil  lll.'i  to  I  l.'i  iiiiii.,  in  llic  lalihil  Ironi  !•!)  lo  KT)  nun.,  in 
the  slufp  alioiil    1.">II  nun.,  in  llic  Imtsc  alHiiil  •JUd  iniii.,  ainl  i an  sc 


'■"■■         ■'  M'''HH       lll..l:..|'   ,  I,  I       .,^.1       -,;::, ,|i      ll,.,fi,l,      :iri^ll, >.■,.!       f,.,       M,..,,|„|y       l||,.      ,„,■,„      ., ,  | . 

;,'"";'  I'"""'  <■<  ■'  i.i'"'i..!.,n  .  .|„  ,,,,„  II,.    Til.   |. ,,--,,!,■  I,, Ml,   iU,  ,>  liii,-,!  »,,i,  .,1111.. ..Ik;!!; 

"""'    '■'"■'    11.. I.   I    l.v    1  I'.mt    „nh    111,-    ,,i.,iM,„,,i,-i     ,Mi.    Ill,     ,.,,,„,il.,    ,-,,r    ili,-    .,ii,n     il    .    f 

";'""''','|   " 'I.  ''"    'I' I'...-   '.I'      i:v    i;.,~  .111.1,,., lit   II   I,   |..--iiii,.   1..   ii.i-h   ..,11   ii,,-   I 

"""'     '1"'""^     ,1,1,  It,  I,  ~     «„|,     ,1„      ,.|.,,;u.,„L        il',..-;,     .1.1, „        l^|.,  nni.Iit.ll      I 'I, , If  Ml,.,  .  ,1.  .>!.■.     ' 

nlicif  hclwciii  llJd  am!  140  inni.  Tlic  iiicssiiif  varies  in  .lill'crcnl  p.i 
(if  111.-  vasi'iilar  system,  lieiiii.'  -.'i  .'ati'st  in  llic  aoila  and  least  in  the  sin., 
est  ailciiojes  lint  llic  fall  in  piissiiii.  the  |iressiirc  ijrailieiit  .|o.s  i 
lieciiine  \c'iy  jiroiioiiiiei'il  until  the  ail.i-iol.'s  liave  heeiniie  so  small  !' 
it   is   no   loii;i(i-   possilile  til  inseil   a   ealinnia   into   lliein;   thus.   Ili 


I  i 


itr.oni)  riii:ssiKi; 


12.'> 


liliMiil  piTssMic  ill  tlic  rcii.-il  oi-  rciiKinil  iiitci'v  is  very  liltii'  less  tlian  tliat 
ill  till'  iHU'ta. 

If  we  cxainiiic  tin  toiiloiir  of  llu  Inttimi  \\liicli  the  pi'i'ssurc  draws,  wc 
shall  liiiil  tliat  it  cxliihits  two  t\pts  dl'  wave,  siiiali  and  Inrfjo;  and  if  wo 
i.lisciM'  the  aiiiinal  wjiili'  llio  1ra<'iiiLr  is  l)t'iii<;  lalicii,  we  shall  (iiid  tliat 


'\'\w    ai1iii.il     III I     |in^-iMr     m..l.!..l     v\illi     a     imrtiiry     man-'iii.lii      ib.u.r     ti.i.im 

.>Mh    a    iia.iiiu    ..I     llii-    TiK|iii.iti>ry    ri..M  m.  iil    ..I    tin     th"i.i\.       N.ili     iImi     IIn     lie  1:11111111.; 
■'■'' llts    ill-lilltlly     Int.. If     lilt      i;^r     HI     !.;.....[     (.it^-iiu. 


I'lnpicf  ai'i'  i-auscd  liy  tin-  licarlln'ats  and  llic  latter  li\  the  rcspii-a- 
-     .Ill   ii|ps('|-\  atiiiii    wliicli    iiiiiiirdiatcly    i-aiscs   the   (|iU'slii>li    as  to   tin' 

'  I'llliiiM'ss  ot"  tlic  inclliod,  for  it  will  In'  .iskcd,  lliiw  can  it  lie  tli;it 
'  '  .iitlicat   ]ii'odiic('s  an  clTcrt    on   Mood   pi'i'ssnrc  wliii-li   is   less  tlian 

'I  till'  respirations.'  (Hivionsly  the  Iraeini,'  must  in'  faulty  in  re- 
I  ti'  the  relative  siyrnilieanee  of  the  waves. 


iL'ti 


Till;  rii{(  ri.\ii(>\  ok    riii;  r.i.iMin 


Spring  Manometer  Tracings 


Tlic  ciiiiNc  lit'  lliis  iii;nTiir;ii-,\    i|('|i(nils  mi   tlic   inert  i;i   nf  the   iii('rciii> , 
:iii  iiiciti.i  \\liicli  is  Ml  liic.Ml  tli.il  I  lie  Niii|i|iii  cltjiiiiri-s  111' lui'ssiiic  j»ri"liii'ii| 
by  ciicli   lic.iilliciil   iiif   licit   iililc   to  o\('ic(iMic   it,   w  licii'.is   till'   imicli    Ic-- 
sitrnilicaiit  Imt  niinc  proloiiLrcd  pressure  eliiiiijies  priiiliieeil  liy  eiieli  res|il 
ratimi  (l<\el(ip  llieir   lull   elTert   on   the  iiiereiny.     These   facts  leil   iiivesti 
(;at<il's  to  seek    lor  iiistiuiiieiits  in   wliieli   the   inertia  error   is  eliiiiiiiateil. 
with  tlic  result  that  they  inveiiteil  what  are  known  as  siiriinj  iikuihiik  h  i ' 


\'lii.    J.I.      lluillili'^    ^J'lm^;    III. m<  tir. 


Many  varieties  of  tliis  instriiiMent  have  lieeii  prodiieed,  liiil  for  our  pur 
pose  it  is  iieeessaiy  to  deserilie  the  principle  of  only  the  simplest  ainl 
most  eflicieiit  the  lliiithle  iiiaiioiiieter.  As  shown  in  I'Mi;.  l!:!,  it  consist^ 
of  a  \ariety  of  taiiilioiir.  which  dilVers  from  the  ordinary  tamlioiir  in  t^^'< 
important  particulars:  (1  i  its  chamlier  is  made  as  small  as  possihle,  aii'l 
(12 1  it  is  covered  not  with  an  elastic  iiiemlirane  hut  with  one  of  leather  or  "I 
thin  Hiited  metal,  'riiese  two  precautions  an'  taken  in  order  to  avoid  spun 
oils  waves  si'i  up  on  account  of  elastic  recoil.  Such  errors  are  furtln 
reduced  liy  liHinn'  the  tuliiiii;  and  cliamlier  of  the  tamlnuir  with  an  ann 
coauulatini.'  Iluid. 


I'll:      -I       .Xit.t.il    I'Mv-ii,     rr...t.!r.|    iiy    a    s|.riiii:    in.iii..iin  t.  i .       'IIk     t  II.  .  l    ..f    \\i:ik    .xmI.iIim' 
tin       v.-ijjM-     i>     V,  ,  n     .'.UTini;      iti.       [.iii'-.l     litilkfl      1.>      llif     >h;m;i1      m.         I  |- imiii      I»iiI...iv,i 

I'.ci'ore    the    tiaciim    taken    with    the    spriiii:    maliometer    can    he    in 
ployeij    lor  i|iiaiit  ilalix  <■   ineasureiiieiits,   it    must    olivioiisly   lie   aradiial! 

a rdinii   to   some   scale      'I'liis    is   accomplished    immediately    lieforc 

after  the  cxpeiiiiicnl    liy  coiiiiectinii   tlic   manometer   throiiL'h   a   T  pi- 
with   a    pressure   Imttle,   which   can    lie   raised   or   lowered    to   a   specin 
lieiirht,  and  with  a  mercury  iiianonieter.     The  displaceiiieiit  of  the  writ  ■ 
point    of  tile   spiiiiy   manometer   coiresponiiiiitr   to   each    !<<   mm.    li'j 
pressiiri'  is  then  written  on  the  traeiii','. 


i!i.<M(i>  i'ui;ssi  i{i 


127 


Ws 


'I'll!'  Iiai'iiius  taken  uilli  Mifli  a  iiiaiKiiin-lcr',  as  shown  in  Vifi.  •2\,  arc 
i|uilf  (lilTfrcnt  rnmi  IImisc  uitli  tin-  niciciifv  nianinnctci'.  It  will  lie  seen 
tliMt  now  the  iMi'iJiar  wavis  aii' dci-iiji'dly  tin'  niiirr  |ininiinni'i'il.  the  i'('S|>ira- 
|cii'\,  liciii";  ciiinpaiativcly  ini'iinspiciiinis.  Instead  id'  thfrc  liciiitr  a  fairly 
steady  prt'ssiirt'  in  tin'  arlciics,  tliis  nndcitrdcs  very  cunsidcialilc  altcra- 
tinii  (luriii<;  each  heartlicat.* 

i'}\aiiiinatinn  of  tliis  ti'acinjr  i;i\rs  ns  accurate  inl'nrinatinn  resrardiii!; 
till'  hlood  pressure  liotli  jpctui'cn  the  heartlieafs  diastrdic,  as  it  is  eailcd — 
.i:iil  diiriii!.'  them  systolic.  It  irives  ns  a  means  of  tcllinfi;  wliat  iniist  I)0 
the  (/((((/  load  (d'  the  circulation  that  is,  the  i)rcssure  that  is  constantly 
[itiscnt     as  well  as  the  lirf  load  that  is  superadded  to  this  !iy  each  hcarl- 


lli;        .:=..         hlltl.llll       1. 1-1.1       (Ml       1  V|.i  IMI.l   lit.      nil       ll"VL.       I.l       -.ll.ru        till        11 1  .mil  1 1  ,  |.|l        •■,        I  111        ^V.|clll.. 

'ii.ivt..lif    .m.l    iiuaii    I.) 1    ).ri--.sim-v    ;it    .litTiniit    p.irl.    nf    tlic    cii.ulat<iiy    .v-tint.      (►    ii    tlu-    Inn- 

■r    /111    prr...iire-.    .iii-l    llir    !,  Mt  r.    t.il.iw    il    i!i.!i..iti     llif    purls    i.l    l!u-    .\>Iimi    t.i    whuh    tlu     inr\ts 
HIT.       iFr.inl    lirul.akii,  I 

l>eat.  This  dilTci-cnce  is  often  calli'ij  llii  in-issiin  piihr,  and  in  man  it 
aiuonnts  to  somewhere  aliont  :'..">  mm.  Ml'.  If  w c  take  tracing's  with  a 
^priMH:  manometer  from  ililTerent  parts  of  the  arterial  tree,  we  sliall  liml 
that,  as  we  travel  towards  the  periphery,  the  pressure  pulse  liccomcs  les.-i 
and  less  marked,  until  tinally  hy  the  time  the  capillaries  are  reached  it 
lias  almost  eutirely  disH|>pe:ire<i.  This  decliin'  in  the  i)rcssuie  pulse  can 
iiiurcovcr  he  seen  to  lii'  dcpenclcnt  more  lai-^'i'ly  on  .-i  fall  in  systolic  than 
11  diastolic  pressure.  In  other  words,  the  dead  load  of  the  circulation 
tlie  diastolic  pressure  rcm.-iins  i)racticall\-  constant  all  alonir  the  arte- 
imI  tree,  whereas  the  systolic  |)ressure  falls  relatively  tpiickly  (Kiir.  li'it. 


■  Hi.  ii.i.  iii„- 
11'    1.111.1  .111.1  t., 


■-■Wll     :ii     I'lk-      .'  ■     1 
■irki'    III   Iht-   I'.rtiii-i . 


1,1    -■4   .111     ii.t    II  |.i..il.    111.     |,iiK.     I  I. 


1..  iiiu    t.'.i    .111. ill    til 


12S 


'iiii;  <ii((i  i.ATiDN  ov  Tin:  m.oon 


Clinical  Measurements 

The   ini'tliiids   (if   lilii(i(l-i)rt'ssiiro   inciisiirciiii'iit    in    iiiiin    Imve   I't'ccii! 
lici'oinc  s(i  pcrl'i'ctt'd  tlmi  the  icsiilts  iii'o  jiliimst  as  iiccuiiitf  iis  those  n 
tniiictl  in  lalmrjilory  atiiiiials  liy  direct  nieasui'enie'it  tlinni^'h  tlie  )ise 
caiiiinhe   iiiserteil   intu   the   ve  seis,      Molli   tlie   systolic   and   the   diastn 
Iir<'ssiire  can  he  nieasnied  with  e(|iial   faeilit\-  and  ai-eui'aey.     Since  tl 
tecliip  •  I'oi'  niakin;.'  the  systolic  nieasui'ements  was  <h'sefihed  at  a  niii' 
earlier  date  than  that    for  tlie  diastolic,   it    has  until   recently  heen   t: 
hahit   with  a  iricat   part   of  the  medical   inofcssion   to  he  satisfied  \\\' 
systolic  readin<;s  alone.    Tins  is  iniist  imfortmiate,  hecause  tlio  kno^^led■J 
which  snch  iiiforniation  irives  us  is  ineoinjiarahly  inferior  to  that  whi- 
i-an  he  ohtamed  hy  <.'an<rintr  the  diastolic  luessure.    Until  we  have  learn^ 
mole  al'ciut    the  dynamics  of  circulation,   it   would   he   jn'ofitlcss   to   •_• 
into  any  details  as  to  the  reasons   for  this  statenuMit,  hut   it   will   sn.i 
hecoiiie  sclf-e\  id   III.     Siif!ice  it  for  the  jneseiit  to  state  that  tin   tlinxtiii 
If'-iin    IS  till   iiiiifi   iiiiiiiirldiit  hiraiisi   it  ijlfis  »,v  tin   loail  ivli iili  tin    /■  ■ 
.Ills  mill  iinrlir  (■<///■/ .v  must  riiiistinitlii  ttutr.  and  the  resistance  wliich  in  . 

I Mfcome  prior  to  the  ()|)eninf:  of  these  valves  at   the  he<;iiiniiiL' 

systol.'.     Moreover,  it  heljis  us  to  <rau>;e  tlie  ix-riphera]   Tcsistanee. 

The  liisi   step  in  the  technic  of  lilood-])rpssurc'  iiicasurenients  in   m:: 
is  the  plaeiiii,'  of  ;in  aiinlet  or  ciifT  ai'oinid  the  arm  or  Ic;;.     This  arin' 
consists  of  a  rnhher  h;iy;  at  least   l"2  cm.  hroad  and  covered  on  its  on' 
surlaee  hy  cloth  or  leather.     The  Ij;i'_'  is  conneet<'d  hy  tiihinsr  with  a  pi' 
sure  <:;iui.'e  ;ind  a  piiiiip.     The  pressure  jiau^'c  may  he  either  an  ordiii,-. 

mercury  manometer  or  m f  the  numerous  (raunes  built  on  the  aiiei'  . 

priinipN'   lli.it    are   now    on   the  market    (  Fiir.   'J(ii.      I-'or   me.-isnrinu;  1 
hloiid  pressure  in  the  vessels  of  the  upper  extremities,  the  arndet  sIcm 
he  applied   around   llie  fleshy   part   of  the  upper  arm  and   for  the  Ihm  • 
liiiihs   around    the   thiu'li.      I'or  accurate    rcadiii!,'   of   lioth    pressures   t;. 
follo\\in<r  jiriiri  iliiri  slioiild  he  followcil.     llaviniT  apjilied  the  armlet.  ''^ 
pulse  should   he  palpate<l   ;it    the   radial   artery,  and   tlie   pressure   in   ' 
armlet  then  r.iised  until  the  pulse  can  im  loin;er  he  felt,  at  which  inoiie 
the  lUessure  in  the  manometer  should  he  noted.    The  cuff  should  tl.ei      ■ 
slmvl\  dec-ompii'sse,|  ,'iiid  the  pressure  noted  at  whicli  the  pulse  ri'appi.i- 
These   t\Mi   re.iiliiiL's   of  s\s|(ilic    pressure   slioiild    he   close    tojrelhcr.    1   : 

thc>y  will  not   iisiiall.v   a'.'r.'c  e\,ictly  I'or  rciisons  which  will  1 xplai   ■■ 

imiiiecliately.     Tln'y  ui\e  us  tin    jiiiljiiilorii  sjtsliilif  liiili.r,  as  it    is  <m'      I 
The  iMcssure  is   now    lowered   .-''.out    l.'t  mm.    Il-r,   iind  a  stethoscdp 

ph I  in  front  of  the  hend  nf  the  elhow   over  the  artery  and  as  clo-      ■ 

to  the  eiitV  as  iiossihle.     With  each  liearlheat  a  distinct  sound  like  ,i  p 
shot  will  he  heard.     The  ileeompicssion  is  now   continued  slowly,  iin  '     ■ 
tlie  pri^ssiire   tails  the  soiimls  will   lie  iieard   to  hecome  louder  and   | 


liT.oul)   lM{i;ssiKi: 


I'Jil 


aliU'  smiii'W  liiit  iiiurii'iliisli  in  qii.ility.  At  ii  cci'taiii  iiicssiirc  tliis  Iciuil 
iliiinii'tcr'  (if  Ihc  soiiikI  will  m|(1(1cii!\  liccipinc  luilcli  less  iiiatkctl,  ;iim1  tin' 
iiiiiiimiiisli  r|M.ilit\-  \(  in-csciit  will  stiddciily  disiiiipciii-.  Tliis  i)uiiit  cur- 
iispdiiils  til  tlic  <li;ist(ili<'  iircNsiiic,  \\liicli  is  iiuw  iciul  <itT  t'rdiii  tin" 
iii:i;iiiiiutrr'. 

It  iini>l  lie  iciiicliilicicd  lluit  liclnw  tliis  jMiint,  iis  tlif  iiicssurc  in  tlic 
ciilT  is  riiilliiT  liiwcri'il,  a  siiiiiiil  is  still  licafd  in  the  aitri-.\  ;  indeed  it 
lines  nut  entiiclv  disaiipcai'  until  llie  iiressiiic  has  lieediiic  i|nite  Iciw.  Tliis 
|ioint  III'  final  disajipeaianee  is,  Imui'ver.  uf  nn  siirnitieaiiee.     The  eulY  is 


'II 


1''!,'.    .''■       Ai'i  .ti;it\!-«    I..r    iri;.-!iiink'    the    .iM.ii.iI    M i    ;ii-:in     in    ii.li:.      Tin-    pif^^nrc    in    Mm- 

irt    l~    r.i-iil    l.y    im-.cUH    ..I     111.'    -Minwi     mini    tin-    pnl-.     .  .m    n.    Iiiiii:,!     Ii.      Iill    .il    ti.v    «M-',       I  hi- 
r-'-iMf    Is    r»a<i    iitT   i>n    tlir    inii.niy    iii..in'iiuui     i~\^niti.     in^^nrt'. 


iHiw  entirely  deei>in|)iessed.  and  slinuld  lie  left  sci  t'oi'  a  niniiient  <il'  inure, 
>'i  tliat  tile  ciieMlatidii  in  tlie  part  ul'  the  aim  lieluw  ii  may  return  tti  the 
iiui'Mial. 

The  aliiive  readin^rs  slnnild  then  he  emil  rolled  liy  a  seemid  iiliserva- 
lieii,  in  which  the  niethuds  emiiliiyed  are  sli'^hlly  nindilied.  With  the 
-litliiisc(i|)|.  at  the  lielid  III'  the  ellinw  tin'  |ilevsiire  ill  the  elllV  is  run  Ul)  1'> 
a  little  aliii\e  the  |)re\i(insly  deteruiiiied  diastulie  pressure,  su  that  the 
siMiiid  is  I'leariy  heard.  I  iie  presume  is  then  lurther  raised  till  the 
siiinid  ilisapjiears.     This  piiint  indii-ates  the  sxsliilie  pressure;  it   is  ealhd 


l:!(l 


I'Ml:   <  IKI  {  l.\'j'ii)\    III'    lili     l:l  mill 


the  ilKililiifi  siisliilii-   iiiih.r.      It    will    lie    t'lillllil    In   \ii\r   il   s.Xstiilir    picssiii 
,1  little  liii/liir  ili.iii  tlijit  iililiiiiii'd  li.\    imlpation  nt'  llic  aftri\   at  tlic  wiisi 
'I'lic  Miiiinl   liiiiiii   iiciw    alisfiit,  tlic    iirt'ssmc   in    tlir  ctllT  is   low  crcd   iiiili 
the  sdiiiid    reapiieais,   aiiil   tlie  ;iiiiiit    at    wliieli    this   (leiMirs  should   allim^' 
exaetly  em  ies|i(ind   to  that    at    wliii'h    the  suiiiid   was   I'liiind   to  disappe,!  i 
II'  thi'  jialpaliii\    N\Ntiilie    index    is   mil    lielnw    the   ainlitnfy.    it    ilidieat' 
tliat  some  eiriif  has  lieeii  made  in  the  applieatinii  of  the  appaiatiis,  aiil 
that  the  readini:  "i"  the  diastolie  inessnre  will   he  liiifelialile.     The  iisu:i' 
siiniee  (if  eirnr  is  in   the   position  ol'  the  >tet  hoseiipe.      If  I'ead.jllstiiieilt    o; 
this  does  not  hrintr  tlie  Iwn  imlii-es  into  proper  lelatioiisliip,  the  aiiseiil 
tatoiy    method   can    not    he    lelied    upon    foi-   eilhef    systolic    oi-    diiistolj. 
readiuL's. 

lu  ease  of  the  failuie  of  the  a useultatoiy   method,  we  iuive  to  fall  ha(  k 
Upon   the  palpatoiy   method    tor  systolie  ))iessure;  and    for  the   measui^- 
men!    of  diastolie,   we  liave  to  use  tlie  inelhod   known   as  the  omllldlnri. 
wliieh   initil   leeeiit   yeais  was  the  on!.\    one   used   for   f;aui:iuir  tlic  dia^ 
tolie  piessufe.     This  I'oiisists  in  ol(seivin<r  the  oseillatiiin  of  tlie  indicat"! 
of  the  pressure  i^auife;  as  the  pressure  in    the  eiilT  falls  irradllnlly   from 
helow   the  svstolie  preN>ure.  these  oseillations  will  he  ohserved  to  inerea-M 
in    amplitude,    until    they    reaeh    a    maximiiiii    heyoiid    wlii<di    with    low. 
pressun    they   lapidly  deeline.     The  pressure  in   the  eiilT  at   the  inoinei  : 
when  the  oseillations  aie  at    the  maximuiii    represents  the  diastolie  pie^ 
sure.     With  a  iiiereiir>    instrument   it  is  ohxiously  ditVieult  to  employ  tin 
method,  hut  with  a  modern  spring  instrument  it  eaii  with  a  little  luaeti.  . 
he  used   with  <;reat   aeeurae.\    and   will  serve  as  a   valliahle  clu'ck  on  tl' 
diastolic  rcadiii;;  as  taken  hy  the  auseultatory  method. 

The  pi-oeediire  may  hi'  altered  in  various  ways,  there  lieiii}r  only  one  [ir. 
eaiilion  to  liear  in  mind:  namejx.  that  the  pressure  in  the  eiitV  should  not  I- 
applied  continuously   t'or  more  than  a   few   momenls  of  time,   for  if  tli 
is  done    for   loiitr   periods,    not    only    will    it    interfere   with    the   aeeura.i 
of  the  readini.'.  hut    il    ma\    e.iUse  eoiisiderahle  diseoml'ort   to  the  |)atieii^ 
There   are   sevei'al    eoiiditiolls   alTeetinu:   llii    iKflirnrif   of   llu     ridiiillilf   li 
each  methoil  whieh  it   is  \\  ell  to  hear  in  mind.     These  have  liccil  iiives! 
LMted  liy   .MaeWilliani.'    Leonard    Mill,-  and    Krlanjrcr.'     With   rcftard   ' 
till  siislohf  /)/•<  vw(//    the  most  impoilant  of  these  are  as  follows:     (  I  ■  Ti 
compression  eiitV  should  he  ;i   wide  one  !  I'J  em.i,  aiid  it  should  never  ' 

applied  so  that  there  is  any  chain f  its  coiiiprcssiuij  the  artery  atrair  ' 

a  holly  surface.     This  preeaiitioii  is  i ssaiy,  since  it  has  hecn  found  tli.  • 

much  less  pressure   is  reipiiicij   to  ohliteratc  any   lierccptihle   pulse  heh., 
the  armlet  when  the  artci\   is  ll.ittened  airaiiist  sonic  hard  striicfiire  th: 
when  if  is  uniformly  compressed  in  the  tissues  in  wliiidi  it  lies.     (2'   l> 
ercpailcies    iire    ofleii    noted    lietnecn    the    systolie    rcadill(I>>    <>ll    conipi ' 


w 


'.I  iM)i>  I'll!  SSI  111: 


Ml 


inll  iillil  i|c('nlil])l('sM(ill  III    llic  alliTV 


I'  III 


thai 


IS,  tlic  |tiilsi'  miiy  ii'ii|ipcai'  <ni 


'Ii  roiiipi'i'ssioii  lit  ii  Idwor  pirssiirc  t'laii  tliat  at  wliicli  it  <liNai)|)i'arril  on 
I'liiiipirssinii.  till'  ilitTort'iUM"  lii'insj:  most  iiiai'kcil  \\  lini  tlio  (Ici'iiiiipn'ssinii 
IS  iloTit'  (inii'kly.  This  (litTciciicc  is  owiiiji  to  the  t'arl  tliat  llif  full  fmci' 
i>i  the  jmlsc  (l(i(>s  nut  icafli  tin'  t'orcarni  until  all  tlic  ncsscIs  liavc  lii'cnni" 
ilistcndi'd  with  IiIoikI,  (.'ii  TIhtc  aii'  nt'li'ii  discrcpaiirii's  in  the  systcilic 
iradincs  takon  t'i'oni  ditTcrrnt  liiiilis;  thus,  it  is  nut  unrniiunnn  to  find 
that  the  systdlic  prcssurr  in  the  lf<r  is  hifilicr  thiin  that  in  tin'  arm  vvt'ii 
iiiit'ii  the  (ibsoi'vcd  pci'snn  is  in  the  hori/ontnl  ))ositiiin.  Thi'sc  ditlVi'iMiccs 
,ii<'  must  eoinniiinly  dbscrvcd  in  ))ati('nts  sulTcriiii.'  fioni  aoilii-  ri'stui<ri- 
taiiiiii  til"  thickciu'd  arteries.  In  aoftie  r-e<;uisritati(in  the  pulse  is  of  the 
uater'haiirmer'  variety,  and  the  ifieatei'  systolic  |)i'essuie  olisei\-ed  in  the 
li'tr  vessels  in  such  eases  seems  to  de|)end  on  diffeieiiees  in  the  i)hys- 
ical  eonditioiis  eoneerned  in  the  transmission  of  this  exaytieiated  pulse 
wave  to  the  vessels  of  the  two  exir-eniities. 

The  I'e.Tson  for'  the  diserepaiieies  in  eases  of  hai'deiied  arteries  is  no 
dciiilil  that  the  lia>*(]eriiii<;  is  likely  to  lie  more  prdiiouneed  in  the  vi's- 
M'ls  of  the  tliifih  than  in  those  of  the  jiriris.  When  a  hai-dened  vessel  is 
iiiiiii)i'essed  it  does  not  eollapse  uiiifoiiiil\  that  is,  it  does  not  lieeome 
iiiiiipletely  elosed  liut  its  \\;ills  come  toirethei-  at  the  middle  part  while 
ehiiiks  still  remain  at  the  sides  The  Mood  eniitiiiues  to  i)ass  throuy:li 
these  ehinks,  and  a  vei'.v  eonsideralily  hiifher  pressiiie  in  the  eiilT  is  re- 
i|iiii('d  to  oliliteiate  them.  That  this  is  ])i-olialily  the  eoir-eet  e\plaiiatioii 
IS  su|)])oi'ted  hy  the  ohseivation  that,  althouuh  in  such  patients  tlie  imlsc 
ihies  not  disappear'  in  the  vessels  of  the  foot  at  the  same  ])ressui'e  as  it 
does  at  the  wr-ist,  a  distinct  chariire  is  nevertheless  ))er'eeptilile  in  the 
pidse  (if  the  foot  ;it  a  cuff  i»r'essure  ei|iial  to  thai  piodueiu!,'  oliliteiatioii 
in  the  wt'ist.  Ill  a  ]»atieiit  showinir  a  systolic  pressure  of  lin  mm.  for-  the 
upiiei'  anil  and  |!IS  mm.  for  the  le;.',  at  IKJ  iiirii.  the  pulse  in  the  le;:. 
allhoiiirh  not  ohlitet'ated.  heeanie  notalily  cut  <lo\\  n  in  volume.  It  there- 
after persisted  at  a  small  volume  with  littli-  alteration  until  the  jii-essure 
Itfcanie  sufticieiit  to  olilitenite  it.  It  is  said  that  lepeate.j  cuiii|U'essioii 
and  dee(inii)re.ssion  of  thi'  hardened  arteries  <«r-eatly  r-educes  the  dis- 
cr'cpancy  in  the  systolic  reailiufis.  DitVcnnces  in  systolic  readings  ai-e 
also  sometimes  ohser'ved  in  normal  individuals,  particularly  after  iiuis- 
ciilar  exercise,  hut  for  these  no  satisfactoi.x-  explanation  can  he  iriven. 

While  palpating  the  radial  artery,  it  will  often  lie  noticed,  as  the 
piessuie  ill  the  culT  is  {ii-adlially  raiseil  from  zero,  that  the  foi'ce  of  the 
jiulso  increases  peree|)til)ly  until  a  pressure  of  ahoul  ."»()  mm.  is  reachi'd. 
ThiK  par-adoxical  l.ehavior  of  tlie  pulse  can  also  he  demonstrated  hy  the 
^I'hyirnio^'raph  I  see  pa-te  -Jlin.  Its  cause  is  not  undeisfo.Mi.  hut  it  is 
•  •f  siffnilieanco  that  the  jrr'catest  auffmeiitations  occur  at  a  culV  i>r'essuic 


V\2 


cimi  t.\Ti(>\  OK  Tin;  iii.iMiK 


I         i 


iit    wliicli  a  Miiiihl   liisl  coiiii's  til  1)1'  lii'Miil  liy  listi'iiiiiir  over  tlic  iirtfi  \ 
at    the  cIImiw. 

\Vitli  ii'iiaiil  til  I  lie  iliiishilli  iirtsntiri,  tlii'ic  liMs  liccn  sdinc  cnntl'iix  cl'v} 
as  t(i  wlicllicr  it  is  iihuh'  acciiratily  iraiitfi'il  !iv  the  (iscillatoi-v  or  tlio  an^ 
ciiltatmy  iiit'tlin<l.  It'lidlli  iiictlinds  ai'i'  ciniilcixcd  it  will  usually  lie  t'liiin.l 
that  tlic  nscillMtniy  -.'ivrs  a  liiul'iT  icadinir  than  tho  aiisciiltatiiry.  Tin 
nmcciisiis  (it'  ii|iiniiiii  srciiis  to  In-  that  the  hittci'  iiictliod  is  the  iiioro  ai'cti 
rate,  ami  certainly  it  is  the  cMsicr  to  apply,  for  with  the  osciljatniv 
there  is  often  <,'i'eat  diflieully  in  (leeidiiit;  Just  exactly  wlien  the  niaxiniMm 
eseillatiiin  occurs. 

Till-  sti'onjiest  eviilence  siippoitiii!,'  the  coiKdusion  that  the  nnseultatm;. 
readinirs  are  more  relialile  than  the  oscillatory  has  Iieen  trained  liy  i'\ 
l)eriinents  with  an  aiti'ii'ial  schema,  consistiritr  of  a  wide  jilass  tnlie  rep 
rcseiitini,'  the  ainilef.  filled  with  Kiiiirer's  solution  ami  closed  liy  i-uM"  i 
stoppers  iiierced  hy  tuhes,  which  are  connected  w  ith  a  fresh  artery,  whiil 
therefore  runs  from  end  to  end  inside  the  tulie.     Tlirouv'h  t\iliin^r  eonnecti  1 
with  the  artery  a  pidsatilc  How  of  oxy^renaled  I»'insrer's  sohition  is  maih 
to  How  at  varyins,'  pressures,  which  are  indicated  by  valvcd  manonntei- 
(see    paj.'!'    I.VJi    connci-led    with    the   artery    fuliin-.r  .just    beyond    the   cmn 
pression  tube.     The  pressure  in  the  latter  is  also  measured  by  a  manmii 
cter,  and  it   is  caused  to  vary  by  a  suitalile  compressor.     I'y  coinpariiiL' 
the   behavior  of  the   artery    witii   the   iiul.satinfr   movement   of  a   spriiii; 
maimnn'ter   connci'ted    with    the   compression    chamber,    tinder   ditTen  n' 
deiriees  of  jiressure  inside  and  outside  the  artery,  it   has  been  observe.! 
that   the  maximal   oscillation   occurs   when   the  artery   is  actually   soni. 
what    tl;ittened  between  the  jiidse  beats;  th;it    is.   it   occurs  at   an  outsiih 
pressure  above  the  diastolic  pressure,  at  w  Inch  of  course  the  vessel  shonM 
retain  its  circular  shape.     When  a  stethoscope  is  apjilicd  to  the  tube  Icailin.' 
fi'iiin   the  ar-triy  just    beyond    ll;.    comprrssion   chamber,    in   the  above  il.' 
scribed  model  sounds  siiiiiliir  to  iIkisc  in  ihe  arm  are  hearil  with  each  pu' 
salion.     While  ijic  pressuie  is  Icin;:  gradually  lowered   from  above  the  nb 

literation  point,  these  sounds  will  be  found  to  1 me  first  audible  as  mmh 

as  a  certain  arm   o  I  of  lliiid  is  for 1  throM!.'h  the  coin|iresscd  area  at  c.-h' 

imlsc   (the  s\st(i|ii-  index  I.  and  to  bccoire  louder  and  often  murmurish   ' 
ipiality  as  the  decoiiipi'ession  is  proceeded  with,  luitil  a  pressure  is  reach'  ! 

at  which  tl'i'V  suddenly  I; mc  less  intense  and  ehanj.'e  in  character.     .\' 

this  moment    it   will  be  ols.rved  by  watchin<_'  the  artery  that  the  extern   ' 
pi'cssurr  is  no  lon'.'i-r  capable  of  pi'oducini.'  any  llatteniiiir  of  the  m's-.  ' 
between  jiulses.     ]]\  identiv,  therefore,  the  chantre  of  soinid  rorrcspon,! 
e\aell\    to  the  diastolic  pressure. 

AVith  ri  liard  to  the  cause  of  the  soinid,  it  should  be  clearly  underst ' 

that    it   is  the  systolic  wave  that  pi'oduees  it.  but   its  occurrence  auil   i' - 


f 


liiudii  i'iii>sri!i. 


i;i:t 


ili!ir;icl('r  wlini    incsciit   Jiic  tlcpfiiilrnl    ii|miii   tin'   iiil  i;i  iiitfiiiil    |)rt'ssiii'o 

(•NistillLT    (llllill'.'    till'    llillsllilil-     pluiM'.        'rllC    I'illlC    111"    llll'    Mllllnl     llilS    lll'CII 

sli'iw  11  to  i|c|icii(l  llll  tlic  |iii-iliictiiiii  III"  M  \\  iilrl'-liilliililt'i-  in  tlir  lilnnil  ves- 
sels lieliiw  the  eiMii|iiessiiiii  eiilT  i  KrliiiiL'er  '.  !>>  a  w  jitel'-liiiiiiiiier  is 
llie.lllt    tile   |ire>Mire  ell;lllt'e^   xvliieii   iire  eiiliseii    liv   sUiliielijy   st iippillfj  tile 

(iiiw  111'  Milter  ill  il   pipe.      Wliell  JI  slldileii   |iirssi||( ills  ill   Vessels  with 

elastic  walls,  tliese  ualls  are  tliiowii  ilitn  \  iliratimi  ami  so  pnxjiiee  a 
SDiiiiil  III  till'  LiUiliir  111'  liliJiiil  pressuie  iiieasllleiiieiits,  as  iiliove  de- 
seiilieij.  when   the   pnssiiic  in   the  elllT  is   lielwceii   systnlie  ami  iliastiilie, 

the  viililiiie  III'  tilt iiipiissed  aiterv    \\ill   increase   nliniptly   with   each 

lieaitlieat  aini  th'is  |ieiiiiit  a  cunsiileialile  vnliiiiie  ot"  sw  ift-llnwini;  lilixitl 
to  eiitee  the  rest  ot'  the  artery  iiinli  nn  nlli  the  eiitT.  When  tiiis  ipliekly 
iiioviiii;  eiiliiiiin  111'  IiIikmI  eonies  into  contact  with  the  stationary  Mood 
lillinij  the  iiiieiiiiipiessed  artery  In  Imr  the  eiilT,  it  will  lieeoine  iiiiinedi- 
ately  cheeked,  and  thus  ili>teiiil  the  arterial  wall  uitli  iiiilisiial  viojeiiee 
and  set  it  into  \iliratioii. 


?l 


I  IIAI'TKU   XVI 

TIM':  r.\(  ToU'S  (((NrKKM;!)  |\  MAINTAININd  TIIK 
I'.I.MOD  I'UKSSI  1,'K 

IliniiiL'  l..'1-cpfiii'  riiriiili^ir'  uilli  ||:r  |ir'iric'i|ili's  .,1'  the  iiii-tliii<|>  liv  wliidi 
''' I   I'lcsMin.    liiciMIITIililils   iilr   liiildi',    llic    next    pi'niili'lll    is   In    rXiilllilli 

ii''"  '1 .iiisis   w  liicli  i>|ii'i'iiti'  111  iiiiiiiiliiin  till-  pn'ssiiic.     Two  "f  iIhm 

caiix'N  nijiy  In'  .•c.iisidcicil  ii>  I'miilfiiiK'iitiil.  >iii('c  willnml  IIm'iii  im  mh-Ii 
picsMilf  ciuild  I'xisI  TliCM-  iili':  ili  tllf  |ilirii|iiMi;  ilctioii  of  tlic  lii-iiil 
.•111(1   (2'    llic   pciiptit'Dil   icsisljiMi'c     Itiiit   is,  llio  ivsisfiiiicp  fit  oiiltlow  <i\' 

'i' 1   frciMi   llu'  immIs  III   llir  ■•iftciiiil  system,      i.css  csstMitinl  tlxiiitrli   im 

piiifiiiit    f;ii-liiis  ;n«':    Cii    tlir    voluiiU'  of  MiiimI    ill    tlic  lilixid    vcssols.    il 

llic    xiscidilv    iir    \  isi'.isit.\    id"   tin-    Id 1,   jiiid    i  ."i  i    llic   rlnsticity    of   llir 

x»!llU  id"  till'  vessels  We  sliiill  Mow  piiieeed  In  exiiiiiille  tile  experiiiieiil.il 
ex  idem-. •  wliiidi  ilidieiiles  Ijir  ieliili\e  iiiipiiiliill I"  eaeli  of  tliese   fjielois 

1.  The  Pumping  Action  of  the  Heart 

< 'liiiiiL:is  pndiieid  ill  tin.  iiie.iii  iiileiiid  liliind  pressure  liy  jilleriitini' 
ill  the  piiiiipiinr  .leliiiM  nf  ill,.  Iiriii't  iire  most  stiikiMi.dy  deiiiniistiiitei]  l.\ 
ol.ser\iiii;  tliis  piessiiie  after  I'litliiiir  or  diiriii;.'  st iiiiulation  nf  liie  vairii^ 

lier\e».  As  will  )m'  1  Nphiilli'd  later  ;  p.iu'e  'JIT',  impulses  eonveved 
tliroip.di   these   iier\es   tn   the   heart    make    the   heals  slower  and    weaker 

Tlies.'    iiiipiils.  s   .irt iislaiilly    arliiiy:    in    the   heart,   so   that    when    Imtli 

xairus    iieiM-s    .-lie    1-llt,    the    heats    In me    ninre     freipleiit     and    stroii'j;el 

witil  the  resiiil  that  the  mean  arterial  pressure  rises  eoiisiderald.w  A 
lesser  de^'ree  of  this  i()V,t  run   Usually    he  nhiained   hy  eiittinir  file  vairii 

nerve  nn  one  side  i  Kij;.  L'7  .  If  l|iA\  the  peripheral  end  of  a  elll  vayii^ 
nerve  is  sliimilaled,  as  hy  apply  inir  an  eleetrie  eiirrent  to  it.  the  heart  will 
either  stop  heatiiii.'  allov'etlier  or  heeoiiie  very  iiiiieh  slowi'd.  with  the  result 
tliat  the  iiieaii  arterial  hlood  inrssiiiT  will  fall,  in  the  former  ease  almost  t  ■ 
zero  and  in  the  latter,  to  .i  le\e|  inrrespondiie^'  to  the  de^'ree  of  slowiiiL- 
of  till-   heart    !  Kij.'.   'JS  ■. , 

2.  The  Peripheral  Resistance 

To  deiiionslrate   the   iiilliiei f   peripher;!!    resistanee   on   ine.'ui   .'!!■!•■ 

rial   hlood   pressure,  the  most    sirikiiiy:  experiment    is  )>eiformed   hy   ein 
tini:  or  stimulalinu  the  Lrieat  spljunhiiie  ner\e       In   this  iieixe  impuKiv 

i:il 


^ 


Ill (•Hl>>l  Ki 


l;t.-i 


-5 


•aIiH'Ii  illT  ••alli'<l  \  ,i-.iiciill>ll  n-lcir  |pcrj|||«.r  lln  \  I'Mii^llirl  till'  llllilfll  III'  llir 
lilmiil  vcsx'Im.  |in>  1 1  allMiiittril  '<>  tllc  lilnml  \<>M'k  hi  t  lir  !lli<l<i|IM'll. 
Tlif  vi'NHrls  nil'  iiinlt'r  tin'  nMNliml  iiilliicini'  I'l'  tliix'  iiii|iiil»iN  mi  (Iwit. 
when  llic  iicrxc-  llmt  liiiiiMiiit  tlii'iii  iiif  srvfii'd,  tlir  msscIs  ililiitc  iiiiil 
lliHs  iitTrr  IrsH  |-t'si>liilici'  In  tin'  liMix  I'llli'llt  nl'  liliiiiil  illmilf  IIm'III.  'I'Ih' 
irsiill  |ii'<><|ui't'i|  nil  till'  ini'iiii  iil'ti  lilll  liliiixl  |iri'ssiii'i'  liy  ciittiii!.'  tlir  two 
>pliiii<-liiiic  iicrxt's  is  llicn-t'iii't'  n  iiiiirkiii  iiinl  ■<iiili|<'ii  tall,  w  liii'li  is  iiii- 
iiii'iliii'  'ly  rrcovi'i'nl  fniiii  it'  tlu'  |M'i'i|>lii'nil  iinl  nt  niif  nt'  tlic  rut  iirrvcs  i'» 
vtiiiiiii.i'ril  iirtilii'iiill.v    '  l-'i^',  L'Mi.     In  rlmnsinu'  llii«  i'\|ii'i'iiMriil  to  |ii'(i\i'  tin- 

!>  Kltinllsliip    lll'tMirll     |li'l'i]lll('l'ill     I'l'sistlllH'r    IIIIll    tllf    lllt'llll    Ill'tl'I'iill     lllnnil 


I'ltC'    ~7.      I-'tltit    nf    cultiliK    llu'    vaKii^    i)t-r\f    iiti    the    atttri.il    M 1    [insMitr. 

|)i'fNMiif.  it  liiilst  III'  ii'iin'inlii'T'i'il  tliiit  it  is  imt  nitiirly  (•(iiit'liisi\  f,  siiu-i' 
the  results  oliseivcij  on  tlii'  iiiciiii  artffi.ii  lilnnil  pressure  riniii  ciitlini; 
nr  stiniulatins;  the  iu'I-m"  in.iv  lie  in  part  ''xplainiMl  as  iliic  to  variation 
in  the  tiititi  ciipacity  nt'  the  ciii-uiat inn  ;  iii  i-  rnoiii  is  created  li\  eutlili^ 
llie   iiciM's,   less   iiioiii   liy    st  iiniilat  mii   tin  tii. 

3.  The  Amount  of  Blood  in  the  Body 

This  ('(III  lie  altered  liy  heiiiniil'.M'.'e  nr  ti::  sfnsinii,  and  tin-  n-Miits 
"t  surh  priieedures  are  (if  interest  imt  iiiily  on  aeeoiiiit  ot'  tlieir  pliysi- 
"l";;i<-  lieariiit',  but  also  lieeaiise  nt   tlieir  ','reat   i>iaelieal   iiiipnrtaiiee. 


1    t 


?  fl 


!i 


'ju^Ji^J 


i.ii; 


I'tll     I  llji  I  IMIHN    ii|.'    I  III     III  )Hil> 


'I'l.  .il'l.r.'fl;!!.'   Ilic  Nii;iiili.iil!.f   (.1    III.'   ivMllN.   it    i>   ilii|i<irt^i  lit    |ii   I.Ciir   li 

IIIIIhI    llllll     '.  .     /.-/„/    ,„/,,„>,     ,,t     II.,     1,1, .,,,1   .■..|i>lltlll.N    frniii    :,    I,,    7    |„.| 1,1 

..I    till'    ^^c•|...||l    ,,(    tlir   ii'iilii.il        Tlil^    tii.'t    li.is    ln'ili    i It'lrl'lli iliiil    pjillly    In 

|M,slll|n||,.||,,     ;|||,|      p.illlv       |,_x      ;|||l,.| 1,1,1      llli'il^lllvlli.-llts.         Ill      tin-     |)llsl 

lll'Tlllll    llntll(Ml.    till'    ImI.iI   ill lilt    111'   IiImimI    i>  ill  tl'llililli'il    l.\    (•..jjirtllm   till' 

'•I '     "'ill''    I'loiilli'-'    till'    illiiliiiil     to    ili'iilli    jiiiil     tlli'll     WilNhiliu    out     th. 

M'ssi'|>   \\iili   siiliiM'   Niijiiiioii    until    till    CM'jiiiiiii.r   ijiiiij    is   no   liiin;ci'   tilini.j 
vMlli   III!.     'I'll!'   Iilooij  I'oiitiiiiii'il   ill   ilir  saliiii'  soliilioii    is  I'sliiiiiitnl    li\ 

i-mIi-Iiimi'IiI"      IIM'IIi.mK      I  M'|.      |M;r,.     "lljl.     ,.|,|i|      is     I|i|i|i'i|      tl>     lllilt      llilll'llv      I'nj 


i 


>i.4M^L 


Sl.->./C',u.b 


~t  _  5t<-i,  ^  Abic 


I'lK. 


i     1 

t       f 


I;m..I      ,.f     .l,i.nil..lM-.K     ih,.     ,,,,|ll.i.il     .11.1     .,f     11,,'     ,i<|il     v.,^,■n^     ..ii     ll"      .,il.n,,l     I,' 

I'll  -sliu. 


''■'■'"■''         '"     ''l''    'I'l''- Iti'lll     Illltlll.ij    sniiil'    vulistill tluit     (JIM'S    not     llil' 

I'uni'    tlirnll-_'ii    M'vsij    wjilU   ,,1-    I,,.,.,,, |uii-k!\     ili'sli'oyrd    is    iliiijcil    to    tli. 

iiioiHJ.     r.y  ill  trriiiiiiiiiir  tin'  riiii.'i'iitr:ilioii  ot"  this  Milist.-iiifc  iii   :i   spiT' 
Ml. "I  of  1. 1. mil,  till'  Mihiiiii'  with  Avliii-li  it   luis  Imciuiii.  mixcil  ciiii  foiiililv  1" 

ciili-uliili'ii.      Ai'.-ii'iii    h.is   ri utl\    Im'cii    I'iuiikI   suiliil.lc    for   tliis   imipn- 

iJtfck',  liiit  till'  l.i'vt  kiiuwii  work  uit'  lljili|;iiu"   w  .is  iloiic  liy  c-insiiif;  tli' 

.■inini.'il  to  iiiN|iii'i'  ji  known  ; nut  of  i-;iil(ou  nioiioxiili'.     This  coniliiiu 

with    till'    li.'iiioL'hiliin    of   till'    liliMiil    isi'i'    |,,i;:i'   4ll|  i    to  ilisiihni'   ;iii    iiju^ii 
qti.'intity  of  o\yj.'i.ii.     j{y  ili'ti'i'iiiiiiiny:  thi'  ililViTi'in'o  lictw  .'cii  the  voIiiiim 


Ill  «H>|I    I'HI  >sl   Id 


1:I7 


.  r  r;il'lM>li   lilMliiiviilc   ill   tlir   IiIimmI    iKl'mr  illnl    I'liHiiw  i|i<_'   it^   ii>lli|jlii<>t  lilt  ion 
w  ,■  r,ni  imIi'iiIjiIc  uilli  liciw    miih'Ii   lildiiil   till'   Kiniwii   iiis|iiii'i|  i|iiMiilily   nl' 

I    lllinll     lllnllliMlll'     IIIIInI     ililVc    fi  iMllpiluc  1         'I'lli'    fCslllls     Mir>      >'  .vllill     III 

illll'i'I'i'tll    atlillli'lN  :    ill    llic    il<i^.    lln'    IiIimmI    cnlist  Itlllo   illmllt     II     |ll'|-    <M'llt 

nl'  till'  limlv  wciirlil,  mill  ill  iiiiiii.  iili'iiil  ."i  |i(r  i-i'iil, 

77i(   itit  till  il  lull   illiit  iif  III  iiinntiiuji   mi  lli>'  M I  |iii>.>,iiii'  i|r|ii'iiils  ini 

the  rail'  111'  lilt'cdiiit'.     IT  a  larifi'  aili-rx ,  Mich  as  iIh'  iVhiiiimI,  is  cut  aiToss, 


^liniiihilinii     of     tlir     Ii-ft     iipl.ituhiiu-     nrrvr     nii     tlu      atti  ti.it     IiIkm.) 
\"\'     ili<      i.rim.iry    nmi    Mwnij.iry    t.^t^. 


till'  |iiissiiit'  will  show  an  iiiiiiiciliat      Mit  iiKnlcratc  fall,  due  laii,'cl\  to  Ilic 


fact  tliat   w  c  lia\c  Miililciilv  tlc( 


the  iicripliciii]   rcsi>taiicc.     1|"  on 


the  other  liruid  only  a  small  artery  or  a  vein  is  ojiciied,  the  iilcediiii.;  will 
at  fust  liriidllee  no  eO'eet  oil  tin'  Mood  |ifes>llie,  and  it  is  only  after  some 
eonsidcrahle  amount  of  hlood  has  hceii  removed  that  it  lie^'iiis  to  fall   ( Fij.'. 


■  !i>i.    To  he  more  exact,  we  ma\  state  that  the  removal  of 


>f  litood  \ii 


kiliiL'ram  of  limlv  v  1'ii.rht  docs  nut  influence  tlie  liKind  prc-isiire.  The  re- 
iiiiival  of  .1  second  iiortioii  of  ,'>  c.c.  per  kiloiiram  causes  the  hlood  jiros- 
Mire  to  hegin  to  fall,  the  fall  of  pressure  for  eacli  sulise<iueiit  f)  c.c.  of 


l:t8 


Tin:    I  IKCI  I.A'I'IO.S    III-'     III!     ISI.IH)I> 


'i    i 


lilnod  \)vv  kilMU'ijiiii  iiimncil  ii\  t'lji-riiiir  alMiiit   t>  iiiiii.   Ilj;,  until  ai'ti-r  "Jn 
I'  IdiMol   per  kil(ii_'i'iim   liiixc  lii'cii   rciiinxcd.   wlicii   n   iiiorc   |-ii|ii.! 


1(1    J-I    IM'.    ( 


III    lniNMirr  M'K   ill    iI).p\mis'i.      Wlirii    flic   proMiiT    rcilrlics   tile   Ic 


I'll  1 1 

"''    •''' -"   to  :!('  mill.    Ilu'.   til.'  iliiiijrci-   Inn  •    is    iriiclicil.    for   tlii-rc    ik.u 

MI|>cr\iMH'N  ;i  Ii-.iiii  i.f  sviiipt s  kiinwii  iis  ••>lnMk."  mikI  till-  clliiliiM-s  I'm-  th, 

.iiiiniiirs  riTiiMTV    Iii'imhiic   iiiicciliiiii.      Tli.-it    llic   i('iii(iv;il   nf  the   lirsl    pur 

''"'I  "''  I'l I-  if  lliis  ir \.il  is  slow   (■ii>«ii>_'li.  lilacs  not   iiillui'iicc  the  It! 1 

pii'sMiii'.  imli. -.Ill's  ilijil  siiMM"  iiil.jiisiiiit'iit  lijis  occuircil  in  tlif  viisciiliii 
s.\si«.ni  In  IkiM  up  111,'  picssiii'c  ill  spiti-  111'  till'  loss  III'  liliiiMJ.  This  itiijiist 
iii'iil  is  liilii'Mij  |(Mui|sivi  in  MisiM-iPiistricliiin. 


.''.WVt-v,'^^ 


Mi,\^  'Av''^ 


i    9- '-novo  I 


L  nd  of 


H     2.  rm  n- 


I  ine  IP  Si.»ci  .» 


fix      I'l      Til.     ilfoi    ,.l    rji.i,;    aii'l    »i"»     h.  iii..irliai!t-    on    tlir    arterial    lilnoil    prnsurr.       Iltlv*.. 
1.1'     ^.l..u,|    ,iiil    lliir.l    |.n-irs    .if    iia.Miit    an    iuKrval    ..(    Iw.)    iiiiiliilcs    i-lai.i.r.l. 

h'l  iiiri  m  frill, I  Ik  iiiiiirl.tnii    is  iciii.'irkniily  nipid,  the  iiri>.'iniil  vcilimit'  m 
hlriMil  liiinir  I'.'siniiil  uitliiii  ji   f,.u    hours.     Thi'  clniiifi's  of  rciovcry  ih 

pcnd  upon  thr  mi nt  of  I>|oim|  lost.     A  loss  cipiiil  to  2  or  :>  \h>v  cent  o: 

the  liodv  xuim|iI  ciii  ,■l||||,,^t  mIw:i,\s  I.c  I'.'.-oM'i'cd  from  in  l.ilioriitor.v  ,iiii 
nulls.  .'Hid  ill  the  I'Msc  uT  nmii  then'  is  rciisoii  to  liclicvc  that  rccoxii' 
nni.v  occur  iittcr  :is  much  .is  :!  per  cent  of  the  Imdv  \vei(r||t  has  lieeii  lost 
The  rcc(i\crv  oi  hjnod  pressure  is  lnoimlit  ahiint  partly  li.v  a  transl'.' 
of  lliiid  rrom  the  tissues  to  tlie  jiloiMJ.     This  .liisti-.-iction  causes  a  dr.vin 

out    of  the   tissli.s,    uhicli    soon   .'Xcilcs  ail   c\treliie   dejili (   thirst.      Tie 

dilution  of  lijoiKJ  li\    iliiid  derived   from   the  tissues  -iirs  very  rajiidl.^ 

as  can  he  sIh.h  n  l>v  I'oiiipai  isoii  ,,f  the  lieiiio>;|ii|)in  content,  or  tile  iiuiiihe 
of    Mood    eorpii^cl.s,    ii,    -..uiiiil,  s    ,,f    1,1 1    |-ci,|.,\e,|    iiiiiiiedi;itel\     hefoi. 


III.IH»1>    I-KKSSIKK 


139 


;in(i  iiniiu'iliatcly  nftfi-  a  li«'in(irihH!.'t'.  The  siu'citic  uiavitv  of  tin-  post- 
li.iiioiThatrii'  blood  is  also  (IccnUtllv  Ix-low  noniial,  iiulicatinR  tliat  tin- 
(liliitiiiir  lliiid  contains  a  lower  coiiccnliation  of  dissolvod  siihsfauccs  tlian 

the  111 1  i)lasina.    The  dilution  of  the  blood  is  indeed  often  so  jrreat  that 

lieniolysis  oeeurs.  the  ))lasniM  Iteinsr  distinctly  tinted  red. 

Ileniorrbniie  also  slitjbtly  raises  the  bydrojren-ion  concentration  of  tbe 
1pIo(k1  plasma,  and  diniinisbes  tbe  store  of  reserve  alkali,  so  that  tbe  ad- 
.lition  of  a  certain  aniotnit  of  acid  to  tbe  libxid  (e.jr.,  carbon  (lioxi<b') 
ijiiises  a  (jreater  rise  in  tbe  bydron;en-ion  eoneentration. 

The  deficiency  in  tbe  blood  elements  produced  by  the  dilution  is  rceti- 
licd  by  the  niamifacture  of  new  corpuscles  in  the  bone  marrow,  etc.,  but 
this  process  in  a  liberally  fed  animal  takes  several  days  f<n-  accoi;ii)lish- 
NM  lit,  and  while  it  is  ijoiiij.'  on  niicroscopic  examination  of  the  blood  will 
reveal  the  presence  of  immature  corp\iseles. 

Careful  studies  of  blood  reireneraf ion  followiiii:  the  removal  on  two 
viu'cessivc  days,  of  I'l  per  cent  of  the  blood,  by  \Vbii)ple  ami  Hooper, 
li.ivc  shown  that  even  in  starviiiir  animals  the  total  anioinit  of  hemo- 
globin (percentajrc  of  bcmoiilobin  multiplied  by  the  volume  of  bloods 
^iiiwly  recovers.  Uecovery  is  irreatly  liastened  by  fcediiift  with  llesb  or 
veil  with  jtelatin.  l{emoval  of  the  s|)leen  or  the  establishment  of  a  bili- 
.iiy  tistiila  <loes  not   interfere  with  the  recovery. 

Incidentally  it  will  be  advantaiieous  to  consider  here  the  effects  of 
transfusion.  These  are  very  difTerent  ai-coidiii):  to  the  iiatuie  of  the  lluid 
i>ic|  for  transfusion.  Three  transfusion  tluids  have  been  iiivestiiratcd: 
1  blood  itself,  (2i  physiolosric  saline  solution  isee  paire  i'.'n,  and  i:'.i 
l>hysiolocic  saline  solution  containintf  viscid  substances  such  as  gelatin. 
Till'  effects  are  also  very  ditTcrent  aeeordiii};  to  whether  the  solutions  are 
injected  into  animals  with  normal  blood  pii'ssnie  or  iiilu  those  whose 
liliiod   ju'essinr   has  been   lowered   by    preeediiiy:   lieiiioriba<;e. 

When  blood  is  injected  into  animals  with  iiornnil  l>lotid  jiressure,  it 
vvill  very  soon  cause  the  pressure  to  rise,  and  as  the  injection  is  main- 
tained the  rise  may  continin-  until  the  pressure  is  perhaps  Tilt  per  cent 
"I-  more  above  its  normal  level.  If  the  injection  is  Ions:  eonliiiued.  how - 
f  \c'r,  a  sudden  fall  of  pressure  occ\irs.  on  account  of  eni:oi<_rcment  of  the 
'iirht  side  of  the  lii-ait.  If  the  injei-lion  is  not  pushed  so  far.  the  increased 
lilood  |)ressiire  after  beiii«:  maintained  for  .i  slmii  time  returns  to  its  old 
!.\cl. 

Iiijeetioii  of  saline  into  ii  iiorniiil  .iiiimal.  if  iii.ide  slow  l>  .  h.is  no  elVeet 

It  ,ill  on  the  111 1  pressure;  if  more  rapidlx    iiijecled.  the  pressure   will 

lise  sliirhlly,  but   to  a  much  less  cNtent   than  that   observed  when   blood 

•!^'•||■  is  injected.     .Much  buffer  (|nantilies  of  the  siiline  than  of  the  blood 

III   be  tolciMtcd    lielorc   cardiac   embarrassineiit    eiisurs       After   tlic  dis 


fl 


J 


1^0 


Till;  nun  i  srins   i>k    i  mi;  mmhhi 


tiniiiiiK r    the    s.iliiic    iiijr.-licii,    ih,.    Iiloud    |ii.>siii,.    ri'luriis    vet. 

r;i;)ii|ly  In  il<.  old  I(\<'l.  'I'ln-  hkisI  sliiKJn..'  itmiIi  •,(  siird  <\|H'iirMiMits  i- 
llii'  (■niniiM,ii>  Miliiiii.'  <i|'  siliiir  s.iliitioii  whicli  ciiii  l>c  sinu  ly  iiijccli.i 
willinll!  |inc,|ilil.l>  JllT.-clili..'  111,.  |.ir.sMiii..  Tlic  i|ll.'sti<MI  is,  \Vlin-(>  .In.  - 
llii'  lliiid  U.I  '  \i  ilir  uriii;ir\  unllIdH  i^  rMirriiiii  il,  .-i  .-nt.-iiii  iiiiTciisc  uii' 
l|sii;ill.\  III'  ,,liN,-|\r.|.  Iiilt  llcv.'f  I'V  ,;ii\  iiiriills  sntlici.'ilt  to  Jli-counl  (..• 
IIk'  (lis,i|.|.c,ii;iiici>  ot'  tlir.  iiij, ■,•(!■, I  s;,lihr.  If  uc  o|>,'n  tlii'  .'ilMJoiiiiiial  c.-n 
il\.  ui'  >li,i|l  fiml  ili.'il  ;i  coiisiiln-.-ililc  lijiiiMhi.ition  ,,(  ihr  smUmc  into  tin 
|in-itori,.:i|  cavitN  llil^  ocnii  i v,|.  ,''ii,|  llml  llir  liver  is  conspiciMusly  c.lrrn 
atolls.  A  .•(■liain  i|. ■.,'!<■,•  of  cirrna  is  also  usually  cvi.lcnt  in  tin-  tissiir- 
ni'  tlic  cxtlcillitii's. 

Still  mole  iiifncstiiiM  ami  iiii|ioiiaiit.  t'niin  a  |)iaclical  stainlpoinl,  ai- 
tlic  ri'siilts  olilainnl  l,\  injccliM-  the  al.ov,.  sojuliuiis  into  tli.'  aiiiiiial 
ulii.si'  lilooit  i>i-cssiii<'  lias  JM'.'ii  lou<i,.i|  liy  a  |ir '\  imiis  li.niotilia-ri'.  I 
the  Mooij  iciiiovcl  iliiiinu'  till-  li.'iiioiiliai.'<-  is  drlilirn  it.'il  i  see  jiaui'  KH 
and  tlini  i-.inj.rlcd  jm,,  ih,.  animal,  it  will  hriiiir  t^  IiI.hmI  prcssiir.'  al 
most    l,ut   not   <|iiili'  hark   1..  its  oiiuinal    \r\o\,  uhich   will    tli.Mi   lir    ''aitl;, 

\>rll   maintain. 'd.      It',  .m  tl tli.M-  lian.l.  s.ilini'  s..luli..M  instead  .)f  li| I 

IS  inj. •.•!.•. I.  the  i-esiiii-ali,iii  oT  |,!,„,d  piissni-e  wifli  an  amount  «(  saliie 
'''|ii;il  '"  H  ^il  111'  llie  ii.nio\.Ml  liloo.r  uili  ;inioiint  t.>  only  ali.iiit  iIii-.m 
•I'l.i'in-s  niiii,.  ..\tent  to  wliieli  it  liad  t'all.'n.  Tliis  partial  ree.iverr  I-. 
i!nii-.',.\..r.  ni,;iiitaiii.M!  Tor  .mly  a  sh.nt  tiiii.'.  alter  \v  lijeli  tin-  pressiin 
rapi.ll\  I'alls  ne.-irly  tn  ih,.  l,.\el  to  wlii.-li  it  uas  r.'.lii.'c.l  l,y  llie  1i,.h; 
.M'l-Iiau.'. 

'I'liese  olisiTx.-ili.iiis  lais.'  tMo  impoiiaiit  praeti.-al  ipiestioiis:  ,|,  \V1,- 
IS  salin.'  ivIatiMl.v  iiieir.'.-t  i\  e  in  llie  test. Mat  i.m  ..|  pressure.'  an,l  •_' 
Why   .li.es   Ih,.    piissiii,.   thus   r.st.n.'.j   so   .pii.-l<|\     fall   au'.'iiii  .' 

'I  he  aiisw,  r>  to  these  .|iii.stions  IniiiL's  lis  to  ;i  eonsi.|erati.;n  ot'  lhi>  ne\' 

ol    th.'    laetors    .•oiie..rii.M|    in    the    maint.'iian. I'    Ih,.    Iiloo.l    pr,.>Mir. 

namely,  the  \  is.-nsit\   „{'  th..  Ii|.i,,d. 


H 


4.  The  Viscosity  of  the  Blood 

Th.-  inip.irl:ii:.-,.  ,.\  this  ta -t.pr  aris,.s  fr th.'  t'aet  that   fa.-ility  ..f  (I..v 

in    a    tiili..    is    iMM'tsel-     proporti.mal    to    tli.>    \is,.:isit\     ..t'    the    tliii.l    ai-i 
'1""'''.^     pr.i|i.irli,in.d    !u   th.'  .Irivin-.r   pi-.-ssiuv   I,,   uhii.],    it    is  siil,.j,.etei| 
"'■'•    '•^-    '"    ""'   'iiiV.  .1  iM-..    in    jn-.'ssiire    li,.t;M.,.|i    (uo    iMunts    in    th.'    till.. 

"'   <l";i''l'""'    ll M'i'Ut    .if   th..    I, .'art    sh, ,1,1,1    r.^main    .^instant,    hut    tl.. 

vise.isit;,    of  tl„.  1,1 1  1„.  d, .,.,■,.,., s... I  |,y  ;,  :„.,||n,'  in.|.M-t i.,n.  th.>  fai'iiitv  -• 

llou  «,,iili|  l„.  in.-reas,..|  ;,i,.|  th,'  pr-ssiire  <l..erease.|.     This  I'.i.'t  .'an  .'asi' 
h>-  sh.,un  .■xperi!u..ntall;    in  a  m.,.l<.|  l,y  .•ansin-j  .„.,im  soliitn.ns  ,,f  vari- 

'■"' iilialii.ns   t.,   I„.   .Irn.-ii    tlir,.iii;h    a    -jlass    ml,.'   l,y    iH.sins   ,f   a   sin. 

pisl.,n    pump  .|,li\  .ihmj  ,i   ,.i,.|,it    .^"loiint    of   lliii.l    int.,    th..   tiihe   \m 


f 


iii.(ini)  i'iti:ssrRi 


141 


J5 


I'.lih  llliivciriiMlt.  Alllldimli  tlic  (iuIIIdW  fidlli  tlic  llJllToW  011(1  of  till*  tlllii- 
iiiii>t  ii'iiijiiii  coiistiiiit,  tlic  jiii'ssiirc  ill  tho  liiliiiii;  w  ill  vmi'v  in  |)iip])intii>ii 
to  the  viscdsity  of  tlio  «rimi  soliitiini  ( liayliss'.) 

Ti'Miisfcrrinu'  flii'so  rrstilts  to  nn  .iiiiintil  \vli<)S(>  Mood  pressure  lias  licen 
Inwcred  by  lieiiiori-liiifjf.  it  lias  been  i'miiid  that  if  saline  solutions  eon- 
t.iiiiintr  a  siinicient  anionnt  of  ginii  acacia  or  '.'ciatiii  to  make  tlie  viseos- 
il\  aliont  ei|lla!  to  that  of  Mood,  ale  lii.icctcd,  llie  oriiriiial  level  of  iilrod 
firissuic  is  recovered  as  well  as  it  v.  oiild  lie  liad  MoimI  itself  lieen  iis- 
jiitid.  A  7  per  eent  solution  of  irimi  acacia  almost  fulfills  tlieso  re(|uiie- 
Miciits,  liiit  unfortunately  tliis  solution  coiitaius  a  slijrlitly  u'l-eater  aiiio'int 
of  calcium  tliaii  it  is  safe  to  inject  into  an  animal.  'I'lie  excess  of  calcium 
may,  however,  lie  renniw-d  liy  r-\iietl>-  ueuti'ali/iii'_'  the  eiim  snlutinn  with 
indium  h.vdroxide,  pieu1r:il  led  ln'iiii,'  used  as  .-in  iinlicator.  .Most  of  the 
enlcinm  lieccimes  |iieci|iii;ited  as  phosphate  The  mucila'je  of  the  lliitish 
ri!aniiaci.|.eia.  diliiteil  !i\e  limes  with  \\;ili-i-.  makes  a  7  |m-i-  cent  sulu- 
tiiiii  of  o-iini  .leai'ia.  A  li  jier  cent  soliiliui!  of  '.'clatiii,  after  lieinir  he.-ited 
t'l  lull  (V.  L'ives  a  visensjty  similar  to  that  of  Muod,  luit  on  aeenmit  of 
•he  possilile  presence  of  teljiuus  spoi'es  siieh  siilutioiis  must  i"'  \.  rv  eare- 
i'llly  slerili/.cii  iM'fcirc  iiijeetinii.  nnd  the  process  of  stelili/at  ion  causes 
.1  decrease  in  \iseo>ity.  The  injection  of  :t  i|uantily  of  one  of  the  alio\e 
-..liitions  eipial  to  tiiat  of  lilnnd  lost  liy  :i  hem-irr!  aue  ^vill  usuan,\  lii'in>r 
the  Mood  pressure  hack  to  its  nritrinal  heiirhi  and  hold  it  there  for  an 
iieiir  or  so, 

\']sciisity  is.  ho\ve\cr,  not  the  onl.\'  propei-ty  of  such  sol''ions  upmi 
v\hieh  their  dcsiralile  elTcct  depends.  The  nsiiinin-  j  r« sxtirr  nf  tin  colhiiils 
ri'so  comes  into  iila.v.  \\\  injectiiMj  s;iliiie  solntii'n  eontaininir  a  siiflicii'iit 
.^iiioiint  of  ;i  eolhiid  siieli  ;is  soliiiile  starch,  which  ^ives  it  the  eorreet 
\isciis|ty  hut  has  no  osuiiitic  pressure,  the  Mnnd  pressure,  allhoiiLrh  it 
t'lNpiir.-irily  recoxeis  ;if|er  t  r;i  Msl'usii.ii,  doi's  not  maintain  its  recovery  in 
tli^  sill  lie  ua.\'  as  w  itii  sol  lit  ions  (•(nitainin'.r  Lrniii  or  ireL-it  m.  The  dilTereiiei> 
liet\Meii  ;|  sImicIi  sohitioii  niiil  iiiic  of  eiiiii  oi'  '.'clatiii  is  tluit  tlic  lofiiier 
i.as  no  (ismotic  lui'ssure.  tin'  ciTeet  ol'  which  is  di'veioped  mainly  on  the 
'  \eietiiiii  of  urine,  as  can  he  sh,>\\  n  hy  ohserviiit;  the  outflow  from  tlo' 
nrelirs  dnrinir  the   in.ieetion    iiit.i   animals  nf  ei|nal   quantities   of  saliiu' 

■ii"ii ■  of  saline  eonlainiin.'  si,ii-.-|i   or  e|.|;iiii,      Knowlfot:'.        With    the 

lifsf  t\;o  thiiils  diuresis  is  produced,  hut  not  \Mth  irelatinoiis  solutions. 
I  he  reas.iii  that  the  osmotic  pii's^uie  of  .ert;iin  colloids  prevents  ]iiiss,-|-_'e 
ot  water  ri-,im  llic  hiood  into  the  iirinifetmis  tiilinies  is  that  th.'  develop- 
iiieiit  ,,l'  ilhs  pressure  mi  the  lilond  sid..  ,,r  the  leii.ii  epil  he!  i  inn  fends  to 
connfciacf  the  liltration  jiressure  by  w  hi.-li  the  urine  is  fornied  ;  see 
!':i'-'e  ."it  I.  I 

Althom.di  the  urinary   fncloi'  will   n,,!   in   itself  explain  the  cffi.-iency  of 


If.' 


'INI     c  li:rrr,\Tli>V    (IK 


II       111  IN  II  > 


"1 


llic  ciilldids  ill  rcciixciiiij;  tilt'  lilimd  incsMirc.  tlic  ciiiKlit ions  ctiiitrolliiiu 
it   reveal   tlie  iiieelumisiii  by  wliicli  the  piissajre  of  lhii(I   from  the  IiUmmI 
vesNels  into  tlie  tissues  is  pii'veiiteil  wlieii   solutions  of  eorre<'t   eoniposi 
tioii  are  iiijeeted.     \oiiiiall.\    tlie  protein  rontent  of  tlie  lilood  ))lasiua  i- 
lii<.'liei'  than  that  nt'  tlie  tissue  lyiiipli,  so  that  there  is  a  <-ontinual  attra< 
tinii  of  water  from  the  tissues  to  the  Mood     an  attraetion  wliieh  is  iii>i 
uiallv  lialaneed  liv  liltiation  t;oin>;  in  the  opposite  dii'i'elion.      When   th' 
fdtl'Hfion    pi'essiire    in    the    Mood    vessels   exceeds   the    ditTerenee    existin- 
lietween   the   oMiioiic    pressure  of  their  eonteiits   and    that    of   the   tissii. 
fluids,  watei-  will  jiass  into  the  tissue  spaees.     When  the  Mood  is  dilutfi! 
as  liy  the  iii.jertion  of  saline  solution,  the  osnnitie  pressure  of  the  eolloid 
in  a  '.'iveii  volume  Iteeoiiies  lowered  and.  the  tiltration  pressure  I'emainin 
e  instant,  fluid  passes  into  the  tissue  spaiMs      (  l|  course  these  ex|)lanatiiii 
rest    on   til''  assumption   that    the  wall   of  the   lilnod  vessels  consists  ol 
memlirane  wiiich  is  perniealile  to  crxstalloids  luiI    impermeaMe  or  neaii 
so  to  colloids. 

Anotlier  im])ortaiit   property  of  the  transfused  saline  solution   to  cm 
sider  is  its  lnnlrDiii  n-lmi  niitit  iilrnlioii.     This  value  increases  in  the  Mcxi 
left  in  the  liodx   after  hemoriliatri',  and  injection  of  sodium  chloride  sol', 
tioii  ay:L'r,i\  ates  tin'  acidosis;  addition   of  NaH<'i>    so  as  to   make  a  •' .' 
M   solution    ri'stoies   the   corrci't    I',,,   and   at    the   same   time   reston>s   tL. 
lost  hulTer  inllucnce  Milioy-         These  ohservat ions  are  of  inter»-st   in  ll- 
lit'lit  of  tlic  icieiit   disco\i'r\   I'l'  t'aiiiion  that  a  condition  of  acidosis,  ,i 
.judtred   liy   the  ( '<  •    comliinins;  jiower  of  the   lijotMl.    is   present    in   shoii 
and  that  the  cle\clopment  of  tliis  condition  can  often  he  <j-uar(h'd  airain 
li,\    liicar'    111, lie   injei'tioiis 

5.  Elasticity  of  Vessel  Walls 

The  clasiicii\    (if  the  \  cssci  walls  is  rsseiilial  to  the  maintenance  of  tl 
diastolic   prcssuic.      If  the   w  nils  presented    no  elasticity  liut    x\  ere   rici  ! 
lilood  pressure  would  fall  to  /cin  lietween  the  hcartlients.     This  fact  c. 
very  readil,\    he  shown  \>\    a  simple  pliysical   model  consistinsr  of  a  pun 
to  icpreseiil   till'  lic.'iit.  i-onneclcd   thidU>rh  a   T  piece  with   two  tiilies,  oi 

(it      Wlllt-li      ,^     i'):lsl)i'        tll<>     iit)i*>l<      IMriil  'l*lii>      I  I'fi*     itiiil      lit'     ii'ii'ti      tiiliii      ll.;      I'l 


^f,       The    free  end   of  c.ich    tulie    is  c. 

tractcd   lo  ,1   iiaitow    apetturc  i  cpicsciitiln;  the  peripheral   resistance,  a 

1...        1...*       .iV    .• .1 1 1'  *      ..  I.     . 


lliHI'M      Ml     ll      liilllim       <l|ICIIII|l-      M-|H  l-scM  1  I  ||w       III,,      f  lt-[    ||niCI  (I  I      resist  illM    I",      il' 

either  tiilir  iii,i\  lie  shut  olV  from  the  pump  li\  means  of  ;i  stopcock  is. 
I'i(r.  IKI  I'^ach  tiilie  should  also  lie  enniicctcd  \>itli  a  mercury  maiin' 
eter.  ll  now  the  stopcocks  ale  aiianv:ed  so  that  the  thiid  passes  ii  ■ 
tlie  riuid  tiilie  while  the  pump  is  in  action,  it  will  he  found  that  w  i' 
each  stroke  ni  the  pump  the  pressure  in  the  tiilie  rises  cimsideialily.  Ii 
that  it  falls  III  zero  lietween  tic  strokes  If  imw  the  stopcocks  are  turn' 
So  that  the  llow   is  thiiiii'.'h  the  elastic  tnlie,  the  .ntion  of  the  pump  \«-'r.. 


A 


Ill  mill   iiii  >si  lir 


1i:i 


iiiaiiu  liilf  Ut'pl  up,  il  «ill  1"'  r<niiicl  lliiil  llir  piosiirc  liclwci'ii  tlii>  strnlirs 
N  iiiiiiiitniiii'it  iit  il  lii'i<.'lil  wliirli  i.s  ilcpt'iiili'iit  (III :  I  I)  tin-  liiti-  .-it  wlni-li 
lie  piiiiip  is  (ipi'i'iitinu;,  iiiitl  i 'J  i  llir  rcsistaiift'  tn  nutfldw  I'loin  tlic  tiilii'. 
rill'  i|Uick<'t-  till'  Mi-tinii  iif  till-  pllllip  and  tile  lii<;lli'l'  tlic  ri'sjstailrc.  til<> 
lUi  r  llic  I'all  111'  pii'sMiic  lii'lwccii  llic  lii'als. 

Tlic  i)liysi('al  cvplanalMHi  "I"  lliis  ii'miH  is  i-li'ailv  tlial  tiic  lliiiil  wittiiii 
Mr  I'lastii-  tulic  whi'ii  Ilif  wave  nt'  pio^iiii'  tiaxcis  intu  it  fiiiiii  tlic  piltiip 
ili^Iciiils  llic  ualU  (if  llic  tiilic.  s(i  llial   w  licii  tlic  |ii  i-Miic  t'rciiii  tlic  pump 


til  act.  the  sliciclicij   chiNtic    wall- 


Ill    III) Iiliiin    III'  Iliiiil 


.Mill  iiiaiiitaiii   the   pressure       \Vc  may    sa\    that    the  clastic   tilicis   in   the 
•  ■--I'l  walls  sTiire  up  xunie  iif  the  -.ystnlic  pics-ii'c-  ami  then  tlaii-iiiit   it   tn 
llii    UIimm!  (iiiriii'j'  (liastiilc. 


!  ■-.;       '<]         lli.iK-im    <1     i  >  i  iritiu  lit     Ii.    -li    v.     l!,.il     tin     -b.i.to;!,      |.tf^-iin      .U  ].,  ii-K    ..rt     tin-    .!.i>ti.if\ 

•  ..     >,...]     M.ill.       Th.      i  ;i|..      11 Inc.. I     l.\     ...ini.T,-Mi.K     ill,      Lull.    (1.     .ii-.ii.|..  ,.i  ~     »li.n     H.ii.l 

..  .     tlir.)ii){li     .111    tl.istl.      t  :'i      ./-')     wllit.     tl.tl.      1^     It  «i.tl.iii.  t'     I ./ I     1..     tin-     i.'ltri'.u         .).     I..i«iii     ..I 
.■...    /(,    I.mII.    Nvnnij.-;    (     ..r-.|    I:.    .l..|.. ...;,, .    /',    ;  mi.l    li.i.,-;    /-',    .-I.i.ti.     ml..:    l.      l-,iM.    I.ll.-.l    witli 


Tli'N iisiilcratiuiis  wniilil  Icail  us  to  c\pcct   lh.it   patients  with  lianl- 

•  iii'd  arteries  shiiiiM  exliiliit   a   lnwcr  iliastnlii-  pn-— .111 ,.  than   iinriiial  jicr- 
~"ii-.   wliieli.   hiiwcxer.   i>.   init    iisuallx    the   case,   since   such   patients  alsn 

-  ilTcr  Iriini  an  increase  in  the  resistance  tu  the  (lnw  nt'  M 1  in  the  peripli- 

' '■'  'riic  (iressurc  imlse  in  these  patients  is.  Iniwcxcr,  \  ci\  iiiarke  1 
"11  the  other  lialiil.  when  the  \essel  walls  liccuine  liiurc  e\!eiisilile  anil 
■  l;istic,  as  ill  certain  cases  nt'  aiieuii>iii,  the  piessiiie  pulse  in  the  \csscls 
l.cliiw  the  aneurism  is  ijistinetlv  less  than  that  iiliscr\|.i|  in  nnriiial  \  es 
M-ls  lit'  the  vaiiic  patient. 


3 
1 


s 


11 


(  ii.\i'ti:k  xvii 

TIIK  ACTION-  or  TIIK  IIK.MJT 

ll;i\  iiiir  stiiilicil  iIh'  iiic'thi>(l>i  fnr  iiii-iisiirciiiriit  .-iiul  tin-  iii.iiii  fiii-lois  i-.iii 

riTllccl    ill    till'    IIUlilltt'llMlll t'   111'   llltiri;ll    lilliilll    lili'sMiri'.    Wl-    IllilV    IIIIW    |i|. 

I- 1  til  >tuil\'  in  iri'i'iitcr  i|i't:iil  tlir  two  iiiii''l  iiii|iiirtiiMl  ol"  llrsr ;  iiiiiiicly,  ll  . 

••iclicin  dl'  till'  lii'.iit,  anil  tin-  ])cii)>ti('riil  poistjinri". 

Till'  lir.iit   Mi-tion  li.'w  til  tic  stiiil'i'i]   t'riiiii  two  \  icw  piiiiils.   lln<  iilnisii.t' 
ailil    till'    lihiisiiiliKliriil.       Friilll    IIm'    plix  sii'.ll     \  ii".\  piiillt    we    liavr    tn    slllil 

till'  lii'ait  as  till'  pump  uf  llir  ciii-iilatinn.     We  nuisl  sec  Imw  it  acts  sn  .i- 

tn  raisi'  till'  ]tri".snii'  of  tl;i'  1>I I  witliin  it,  ainl  Imw  tin-  vahcs  npi  ra'' 

sii  as  til  liiiri't  llic  lilunillliiu   alwaxs  in  mii'  iliri'i'l  imi.     We  must  also  i'\ 
plain  tin-  ciiusis  uf  ci'itain  si'i-umlai-.v    pli.\sii-al   iilirnuiinna,  sncli   as  i\,-- 

lirail  sduiiils  wliii-Ii  ai'i'unipany  tlif  lirart  ai-tiiui,  an. I  ut' i-i'ltaiii  si iiil.ii^ 

cliaini.'s  in  |iri's>uii'  prmlui-i'il  in  tlir  ntliir  llimai'ii'  \  isnTa  liy  rarli  Ihm'' 
liial.     I'riiin  tlic  pli.\  siiiliii.'ii'  \  irw  |iiiiiit  \M'  must  iii\  rstiijati'  tiic  I'mnliti"' 

iispiiiisililr  I'nr  till'  riiii>taiit   rlivtlimii'  artivity  nl'  tlu'  lirail   ainl  tl < 

tinl  111  \xhii'li  tliis  is  Mili.ji'i-liil  Iliiiiuijii  till'  lU'iviiiis  system. 


"\ 


THE  PUMPING  ACTION  OF  THE  HEART 

Wlii'ii  till'  lirart   is  \ii'\vi'il  in  tin'  npnn'il  tlmrax  of  an  animal  kept  ai  \ 
I.,    arlilii'ial   ivspiratiuii   ami   l\in^  in   tlie   pnnii'   pn^iliun.   it   i-an  I"'   ii"'     ' 

ill  at  \\  itli  lai'li  i-iintrai-ticiii  tlie  xiiit  lii-lrs  In me  ^inailii-  ami  lianle'-,  i  i 

the  api'N  teiiiK  to  rise  up  a  little,  sn  tlial  if  llie  tliiir,i\  were  iiitait 
wiiiiM  press  mule  firmly  an;iinst  tlie  walls,  ainl  tli.-it  it  rotates  --W'J 
I'rom  lel't  to  riv'ht,  lillt  i\nv.  not  inii\e  hearer  tlie  liase  of  tlie  lie.irl,  I 
till'  aiirieuloxciitrieular  <;roo\e  is  eaiefully  o!i-.ei'\  eil.  it  will  often 
lioleij  that  it  moves  sli',;litl\  lowalil  tlie  ;ipe\  with  eaell  s.vstole,  \\1mi 
till'  liase  of  the  heart  it>-r!f,  \' hne  it  is  altaelie'l  to  the  lar'_M'  vessels. 
mains  tlxed.  The  aiirieles  eail  often  he  seen  111  eoiilrael  ainl  relax  l" 
the   \  elit  rieli  s. 

The   most    iiote\' oithy    results   i,l'  this   iiispeelion    are   tli.it    iluiiii'.^ 

toll-    the    ;ipe\     of    the     jii'art     ilors     not     liioM      ti'Wal'l     the     liase,     lilll     ■ 

the   am  ieiiln\  enl  I  ieiilar   tiroiive    iiioms   sliuhllv     tow.iil    the    apex       'I 
these  same   iiio\  i  lainl  s  oeeiii    in   the   iiitaet    :,'imal   eaii    he   shown    i'\ 

\eiy      simple     (Apel  ille   lit      "f     pllshiliU:      ■■    o      liillU      steel      Kllittill',r     le-'  >; 

lit 


Tin:  ACTION  or  Tin.  invici 


141 


Illloll 


;li  till-  tliiir.Mfic  Willis  into  tlic  licnrt  walls,  oi f  tlinn  so  ]»lii<'i'(l 

thiit  it  i>i<Tccs  tlic  apex  of  llif  vciitiicli-.  tlic  otlnT  so  tliat  it  picn-cs  the 
Till-  iici'.llcs  tlH'ii  ai't  as  levers  willi  tlicir  I'lili-ia  at  tlip  clicst  wall. 


MSC, 


,iih1  if  tlic  iiiovt'MH'llls  of  thi'il-  outer  free  emls.  pro.liii'e.l  liy  the  iiioveiiieiits 
..r  the  heart,  arc  ohscrved.  tlicy  will  he  found  to  coiiliini  the  oliservalions 
made  DM  the  exposi'd  heart. 

Mine  )i;irtieiilar  invcNtiuatioiis  of  the  chaiiu;es  oeeiiiriii<:  in  the  shape 
u\  the  heart  cavity  diiriiiu'  sy>toli'  and  iliastole  \\n\r  hecii  iiiidi'rtakcii  hy 
m.-ikini;  nicasiiremeiits  of  sci-tions  across  the  heart  in  one  or  other  of 
tliise  conditiciiis.  h'nr  Mich  piiipoMS  the  licait  ill  iliiistole  is  easily  oh- 
t, lined,  hut  for  the  heart  in  sysl.de  it  is  iicicssiry  to  use  tjic  M.iiicwhat 
nililicial    iiicalis  i.f   injcctiii!.'   thi'   heart    uitli    Imt    chnniiic   acid    solution 

just  heroic  the  de.ith  of  the  animal.     Tl hriunic  .mcIiI  caiis's  the  cardiac 

iiiiiM.|,.  t itraci   and  iiiaintaiii>  it   in  this  cniidiliiin.     The  oiitccnio  of 

tlicM'  in\f>tiu'atioiiN  is.  however,  imt    <<\'  iinich    practical    iiiipnrtance. 

.\|tl irh  it   is  iiiiw  c.iiiiiiion   kiinulcdue  that    the  din  d  ion  of  the  How 

i.r  the  hlnnd  is  fiuiii  the  Mills  to  tlic  aitciics.  y.  t    it    may  he  <•(  interest 

\ Msidcr  for  a  moiiieiit  the  uciicial   principle  uf  the  methods  hy  which 

\Villi;;m  llaivev  succerded  ih  makinu-  this  d'se..\ciy.  His  evidciK'c  was 
).:iill\  aiialomic,  partly  experimental,  lie  pniiitcd  out  that  the  wiills  ot 
the  \ciiis.  and  of  the  auricles  to  which  they  lead,  are  very  th'ii.  whereas 
tlinse  lit'  tic  arteries  ami  \ciitiicles  arc  very  thick,  and  he  coneliid.Ml  th.it 
ih  til.  veins  the  hhind  'iillsl  IIhw  -lelitly  Ironi  the  tissues  tow.'ird  the 
li.arl,  tl.  which  the  valves  in  tl:c  veins  diicet  it.  and  that  in  the  arieries 
i!  must  he  i.ropclled  li>  pnUcs  with  e.nh  svstnh'  tliri.iiiih  the  arteries 
Inwards  the  tissues  li\  the  cniit  lad  ii  n  of  the  w.alls  of  the  ventricles.  Tin' 
I  xlicriiiicntal  su)>port  for  this  hyp'Uhes's  he  fiiinishcl  |i,irtly  by  claiiipinir 
the  laisrc  vessels,  veins  and  arteries  le.-iiliiej  to  iir  fiuni  the  heart,  and 
i.liMTvini.'  the  resultinu  disti  iisiiiii  m  cnllapsi-  ot'  the  vessel:  and  partiv    \>y 

lali-iilatinii    of  the    amniinl    of   Itloud    .vhich    niiist    I \pelled    from    the 

v.iitiicles  in  a  irivin  period  of  time 

llaivev's   discdv  cries  ennccfliili'.'  the   (Vclils   nf   the  eardi; ,velc    were 

iml    u h   ailded    to    until    e\pclii tal    liietln.ds    were    dcv  isid    h,v     which 

the  I'lcssure  ehaii'-'cs  (leciirriiiv:  in  the  various  civitics  fnuld  he  ineasiired 
and  i-iimparcd.  I'ntil  sudi  mcasnreuients  v.eii'  el.ilinr.il  id.  il  was  impos. 
sihlc  to  investiirate  the  nieehanisMi  li.\  w  hieii  the  various  valves  l.etueeii 
the    he;irt    cavities    and    the    vessels    eoniiected    with    lllclll    perfiUIll    their 

fillietinn.  nr  to  descrihe  with  ali.v    de'-'i( f  aceur.'ie.v    the  events  ncciiiriliL' 

III  the  he.irt  chaiiilicrs  diiriii!:  the  various  ph.-iscs  nt'  I  hi'  cardiac  e.vele. 
It  is  fur  tiic  jiuiposc  III'  aseertaininy;  the  cx.-ict  lime  relationship  of  these 
•  liaiiu'cs   that    intiaeardiac    pressure    eiirvis   are    studied. 


l»tl 


IMI     (  li;c  I  I   \Tlii\    (1I--    Till     I'.l  iiiih 

Intracardiac  Pressure  Curves 


Till'  t'liilit^t  inc'tlMiil  I'm  t;ikiiiL'  siu-li  ruivi's  coiiNisti'il  in  intl'oilui'iiiL' 
into  tin'  I'iinliiic  I'liiiliiliris  mill  tlir  MimmI  M'sm'Is  111'  llic  liiiisr,  sii-callril 
iiiriliiic  siiiiiiils  I'lii'si'  (•iiiisisli'il  111'  ii  iiiiiii'  or  ll•s^  ii;ri(l  liilic  fiiriiishfil  ,ii 
iiMf  t'lid  with  ii  littli'  ^la^tll'  liai;  or  iiiii|)lillii  ami  I'liiiiiii-lfii  :it  tlif  iillii'i 
\\\t\\  il  tiiiiiliiiiii',  l>\  iiii'aiis  lit'  liililii'i'  tuliiiit:  <hif  III'  tlifsc  littlt'  liau- 
was  placfil  ill  mii'  nt'  tlii'  \riitiirlr>.  aiinllii'i'  in  tlii'  aniifli'  nr  amta,  tli< 

mill'  lii'iiiL'  iiiM  itiil  In  llii'  I'liiiin'r  I'M'^r  t linmi'li  m |'  tlu'  laiut-  \fins  al 

llic    liHit    111'   llir   111  rk  ;    ill   llir   ImIIiI    imm'   I|iIiiI|i;1i   llir   raii'tiil   aitflV        Tin 
iiitrai'Mi'iliiii'    |in'>Miir   iiiims   nlitaini'il    in    this    wav    iiiarki'il   a    nii-at    a<l 

viiiii MT  till'  iiii'tliiMN  that  hail  jircv  iiiiislv    hii  n  usiil  In  stiiilv   the  I'vriii- 

lit'  tin-  caiiliai-  I'M  li-.  hill  tiirx    \Mii'  Ml  I'ailltN    in  iniiipai  ismi  with  tiariiiL' 


.,^ 


---b 


C-- 


"^ 


I'lK       .<-•         |1,..K,.,„.         I      WiKK...'      ..,!.,  ,1      111 in,  I.'  Ih<       «i,,.       nlj.,      nil..-      l.tl       I.    .1111.,). 

"'Ill    'Ii.     Ml  If,.!..    ,1,    •     I- t..|     ».ll,    a    i.r.i,>    ,\lin.|ii     I  K  i    |,i  .,t  i,|,',|     ailh    ..    »l..|...„',     .i 

lilt     himi'ii    ..f     wliuh    »..iiif»    in    .i|>|ii'>itt>>ii     wilh    j    (  l.ilr     iji     I):i\imk    ,t    small    «.|'*III|1K    III     ll.        I 

"•'•l""i    ■■'    ' .■11,., .11 I.,ui,,.,    /(    ,,,„|    a    1-    i,ii;l.ili.|    I.I     111,     |,„sil„,i,    ,,(    111,-    lap.       AI...Vf    a 

■'    "Kiiu"!    <.i|..iil.'    iM     I    Mill,      ill    .li.,iml,i    aii.i    ,..»■, t,. I     l.>     riil.h.r    .l.iiii        Tlii»    larric.    .1    -• 

i.iitr.r     11   I     f,,.|iii,,|    -..    .h..i     II     IH..1 il„     ||,..,.l     ,,.|,     ,,     ,|„.    >,„.,|i|,.         Alii.vc    ill.-    ,.i|.miI, 

.iirjiiB.-.l  .,11  iml,,„,l  11,1, ,,.r.  li..|ii  „l,„|,  .,  -it.,,.,;  !.,  .iiii  .1  IikIii  is  i,H.-ili-.|  ..i.  I,.  111.  Mi.i- 
''■  ""  ""  'il'^iil.  Tills  1.1,1111  111,  I,  1..4,),  l.a.k  ,,„.|  11„.  11,1, ,,.,  ,/i  I,  ;i.||ll-l,.l  -.  ll'..- 
inii-Mieis    ..„     .,     Ml  .>i.,i..     |.li..i.,ui.i,  1  I,      il.ii,  ri„      -1,^.1,!,, I     ,i,.,>,„i,iii,     ,,t     till-     small     liiiii-i 

II.-     lliiis    k-r.  ittlv     !i;.iKti.i  1.1 

taken  hv   iiiiiii.  iiiiuh-rn   in.tlii.iU  ijn.t    it    is   nut    wnith    vvhilc  cimsiiliTii: 
tticiii  aii\    t'ni-tlicr  Ihti- 

'I'lli-   |ih>sii-al    I'lliil-s    ili\i,l\ii|    In    t|||.    usr   ul'   Ihr   nhllT   iiisti-iniirnts   ^\r\ 

ilili'    liiainix    til    thi'    I'hisin-    t-, i|    ,,|'   ij,,.    nn'iiihiaiii's,    i-ti' .    ilscii    in    tin- 

(■(iiistnii'tiiiii      A  jiirat   iiiipi'iiM'iiii'iit   in  tcclinir  was  alTiiiiltil  hv    tin-  u-^ 
lit'  till'  s|ii-ii:^.  iiiaiiiiiiii'Iri' .ll    lliiitlilr   i  si'.'  |ia^'i'   iL'lii.  uhii-li  was  i-miii.— t. 
with  mil'  lit"  thi'  hrait   i'a\itii-s  h\    a  laiinnla   lillnl  hcrm-c  iiiscrtimi  «i' 
sotiii'  ;intii'iiai.'iil;inl  tlniil      'j'lii'  i-av  it>    nf  thr  tainhmir  was  iiiailc  as  sm, 
as  (III     ihh'.  ,'iiii|  i-ithi-r  h'l't  i-iM|it>   111-  filh'il  \vith  thr  antii'iiaKnlut  in'-  Ihn 


Ill      \i    I  Ins    III      I  ill      III    \IM 


117 


A  sciin-liiiii.'  iiiv  I'Nlitriitinii  iiitn  lln'  |i|i\  Nicil  pi  iririplrs  uimiImmI  in  tiiK 
i;  irctiids  of  siidili'ii  cliiiin'c-i  in  |ii(  nmih-  Ii\  >iii'|i  irislniiin'iits  luis.  Imw 
(T.  slioNMi   tliiit   .Minsidfiiilili'   <i  lilts   iiiT    jiiiiiiiTi!,   till'    iii'iliii   <>(  ll" 


'i.Iiiiii;  mill  IIk'  iii'rcssitN    nl'  usinn  Ii-mms  hi 
M'   ii'>i  ip|i>i|i|.-   |ni    III"''!    I'l'  III!  Ml   ii-t'.   W m 


rcrcilllx    Ih'i'II  iii-llirv  til   li\    llsiiiu 


it   ^11  r;i 


lll'.l 


iiiiiv iiitr  iiiii>.>  111'  lliiiil  ill  I' 

■  il-ijcl'    III    si'cuii'    IrriHi' 

'.'ti>).  Tlii'ii-  cliinimit 
..|ilical  iiiiiiiiiiiictn-.  (iiic  III  nliiili  i  WiiJizcrs' '  is  siinu  ii  in  tlif  (ic'imi 
|i;iii.vimr  tiiriiii'  It  imusIsIs  oI'  a  wuiv  irias-  lulic  .1.  riiMiii'''tcil  aliii\f  wiili 
.1  liiilliiw  lulls-  rviilKlcl  /{.  pniviilnl  with  a  st(i|)iiM'k  ('.  ihi'  lllllM'll  of  which 
laliiTs  fliilil  lichn*  liI^Nald  till  it  asMiincs  tin-  saiiH'  iliaiiii'li'i  as  an  apci 
|iiic  ill  the  scjfiiKiil  rajisiilc  h.  almM  il  that  is.  a  .aiistilf  nit  iiw  a>  at  "nf 
mil  wliii'li  is  :i  Mini  in  iliaiiu'lff  anil  i>ivii.il  «itli  nililMr  ilani.  Il.v  ail 
lustiiii'iit  III'  this  s|i>|iiiii'k  till'  piilsiitiiiiis  III    llii    lliiiil   in     I    anil   //  fan   I"- 

■l,IMI|il-il    III    a    Itlrali-l     111     llss    i\li  III    111  liilr    llli\     air    II  , -111-1111111  i|    illtli    ihr 


I  IK  .M  lli'tua!  ffinrtl-.  t'f  iiiIraMiitri^  iiiar  |'U-'iiii;  al.  rimit-iilar  *>'*li.lr;  !•  J  i-ii  ■•i-li>Kiiih 
.MM, I;  ,11,  «|ili\iiMii.  i.rri.iil;  allt  1  (,  .Ii.i>1mIi'.  ni-lniiiriil-  .1  \.irMiiu  ,l.gr.T>  ..l  -i  ti.ilu.ii.  .- 
^iir    rinpliiynl    in    t.tkiiiK    lilt'    iiir\i-       il'ioiii    \ViUil<i^  > 


il 


-••'.Miii'iit  capsnlf  A  small  piece  nf  I'l'llulnid  I'airyinir  a  tiiiv  miii'ir  ifsts 
'III  till'  iiililii'i-  ilaiii.  Iifiiii;  piMitcil  on  thr  i-imril  siilr  of  the  fapsiih'.  A 
iiiirrnr  is  attachcil  tn  tin-  rapsnlc  with  its  phmc  si«  aijjusti'il  that  thr 
niiairc  of  a  sti-nii'^'  liirlit  plai-cil  at  sonii-  ilist;iiii'i'  t'lnin  it  is  t'ocnsnl  on  tin' 
iiltic  niiri-iir  rairicil  l>>  tin'  i-i'llnloiil  Tin'  ia\  nllii-li  il  Iruiii  I  In'  litth' 
iniirnr  ami  airain  iclh  rli'il  fioni  thr  laiuii  niirinr  is  ailjiisttil  sn  as  I  i 
iuipinj;c  upon  a  nio\iii!x  photof^iaphif  plain-  tiaxi'llinir  at  a  iinil'oini  lat  • 
III  a  siiitalily  ronstrin-tcil  phiiloiriaphir  apparatus.  \\\  the  use  of  sin-li 
an  apparatus  tin-  rhii'f  errors  I'lii'ipunti'ieil  li\  Ihr  use  uf  tlie  nldrr  in 
stninn'iits  are  eliminated,  lieiaiise  tluic  is  im  iiio\  iiii:  mass  of  lliiid  and 
ilu'if  are  no  levers  to  set  up  spniious  \  ilnatioiis.  (iiixes  sei-uri'd  li.\ 
'lie  Use  nf  this  iiist  lunieiit  a'e  slmu  n  in   I'i'^    ^{^t. 

Two  olijeets  must  lie  kept  in  view  in  aiialv/.inj,'  the  eiirves:  (li   Curxes 
"litaiiied   from   the  dilTiient    eavities  ma>    lie  eonipaicd   in   order  to  de 
'••rinine  the  exaet   inoment   diirini:  the  laldiae  evile  at    wllieli   stieh   pus- 


n.H 


Till     <  lie  I  I  \  iiiiN    UK    rill     111  iKHi 


MllT  rh.iiit'i'N  ..cciir  IIS   iiiiisl    ••■\\v   III   |.r<ii|ii |i*>iiiiii{  or  ■•IiihImk  of  lli. 

Miii.nis  viilvts.  iiiiil     2     111.'  < luiir  d'  tin'  ciirvt's  nlitiiiiM-il    iVmii  cacli 

..i\itv   iiiii.\    l.r  r\iiiriiii.'(|    in   iti'.lt'i-  t.i   liiiil   out   txii.'llx    hiivv    tlic   IMcsmii. 
ill  flial  pailiriilfir  cavity  is  IhIi    xin-.'. 


Compariion  of  the  Corvei 

Il.'fnif  ii  iiiL'  llic  ciirM^  r,,|    .!><■.  rlaiiiinv'  llir  i'.liifi\i'  pti'ssiiiv  in   tlic 

ililTn.'iil    .•a\iti,.s,   llir.v    M    isl    lir   -vailii.it.'il   ; riiiiii;   tii   Mini.-   scali",    I'l.r 

It   is  rliar  tliat  l.\    tlir  iisc  ,,|'  iiixli  mii.  iiN  \iU,-  ihusc  w,-  hav.'  Iir.-ii  <l.'sri-ili 

IIIL',   lllf   alisnllltl'   pl.'ssmc    \allll'   ..I    .  .irll   CIIIM'   will    \ ,.    \    ;u nlillir   111   til. 

.•..iistiu.'ti.'ii   .il'   til.'    iiisti'iiiiii'iit    1  tjii.kiii-ss   of   iiiciiil.iaii.',   (ti-  1.   ami    in 

.l.'.-.l   invliiiiii,.|i|s  „(  varviiiL'  .l.'ur.  ■ -.  (      icsislj must   lir   •  iiipl..\r.|   in 

lakiii!.'  iMiiv.'s  In. Ill  pla.-.'s  liavi'-j  sii,  h  ,|  ir.'i.nt  pifssincs  as  exist  in 
til.'  ami. -Its  ami  \  i  ni  li.l.s.  Tli.  uradiiatimi  is,  li.iw  cn  cr.  a  v.i.v  (•a>\ 
iii.itl.-i',  an. I  .•..iiHsts.  as  alr.'a.|\  .|.|aiii.',|  (  ;.air.'  rjtii,  in  .•..|iii.-.-l  iii^'  tii.' 
itistiiiiiii'iit  liy  nii'aii->  .,1' a  T  pi.'.'.'  w  il !  ,  iii.'i.'iiry  iiiamniii'tci'  an. I  a  i>r. - 
siii't'  li.,ttl<'  ami  linn  iiiaikiii','  .ni  tli.'  iia.-iii'.'.  tin'  p.iints  <'..in'sp.iii,|in..r  t,, 
.•a.'li   In,  Jii  ..r  :>il  iiii'liin.t.i>  ..f  iii.'i-.as,'  ..l   pi  .'ssiir.',  as  llic  casr  ni,i\    I.. 

'•'"  ■■' '•'•'••I  til'-  tiiiii'  i'.'l.'iti..nsliip  li.'tui-.'ii  III,.  ,,|.,  Mill','  ami  tlii'  .'L'sliii: 

..r  til.'  aiiii.'iil..\t'iitii.'iilal'  Mihc,  tin'  tracings  slnnil.!  .■  takni  IV, m  tl 
li'jhl  ;iiiii.'lr  ami  tin'  li'.'lil  \  .'iil  ri.-li',  .'in.|  t.>  asccilain  llic  sann-  uitli  ■ 
'.Ml. I  t.i  III.'  sriiiilnnai-  \al\.',  Ir.nn  tli.'  lift    \  .'iitiii'lc  am!  tlii'  a..i'la.» 

l!>    .■.Mnp,iriii._'  til.'  ciiiM's   it    is   mm    an   casv    niatt.'i'   t.i   .isi-.Ttain   tli- 

<  Na.'t    III. .III.  lit    at    wlii.'li    .|i,.   pi-,.>Min.   jn    tl n.'   .'jivitv   .-.imii's   t.i   i'.|iial 

'''■''   '"  '' ""'I'      'I'lii^  limnnnt.  r.'a.l  .ni  tl|.'  a.'.'.niipa  n\  iii'_'  time  tra.'iir.', 

"ill  ..l.\i,,iiNlv  imji.-at.'  Ilial  al  uir.-li  llir  pari  i.-nlar  v;ilv.'  is  jiist  al>.>nl  l" 
"I""  "!■  'I"-''  ''"'III  '!"'•  i-.">iilts  .if  su.'h  .'sp.'iiiinnls.  tli.'  i-iirvi's  may  I..' 
Slip.  limp. I-.. '.t  as  in   l-'i'j-    :il. 

Ill   tlic   liist    pla.'.'  j.'t    lis  c.nipan'  tin   ciirris  frimi   Hi,    rli/lil  aiirich   ,(//./ 
r.iilnrl,.     'j'hc  .iirv.s  li.'Lrin   at   tli.-   \rry  .'ml   .,f  .liast.,!.'.  ami   lln-y   sji.iu 
Iliat  ;i  .listiii.'t   iii.i.a-.'  in  pr.-ssiin.  js  .i.'.Miiiim.'  in  li.itli  .'iiiiii'li-  ami  v.-n 
tri.'l.'  ami   lasiii-j  a|„,nt  ii.iCi  s.'.'..n'i      This  is  ,,f  .'.iiiis.'  I'.'nisi'.l  l.y  ami. 
niar  s>Nt.,|.'.   ami   sin,.,'   it    nf.-m  >   in    I...1I1   .-axilifs.   it    imlicatcs   that   th. 
I.as-ai.'.-    |..'t\\.'.!i    tliiin,    tli.'    aiiii."il..\  .nl  ii,-iil,'ii'    ..rili.-.  .    „iiist    l.i'    ..p.  1 
'•''"•   '"'"    ■"!;'•■  '•"'•^'-  'li"ii  -u. l.i. 'Illy  sli.H.ts  auay  li..y..m|   tli,>  auri.'iil 
'"■''""•''■  "'    ''"■  "ii'"''l   "I'  \\-i'.l.'  ill   til.'  v.nlri.'l.',  ami   tin'   p.iiiit   at   wUu 
til.-   lu,,   .-iirv.'s    |i...jiii    I,,   s.parat.'    imli.-at.'s    l|i.'    iii.nn.'iit    jii    nlii.'li    tl.^ 
.'iiiri. '111. .x.'iilii. -Ilia!    MiK.'s   .   MS,'.      {•■r,,in    this   time   mi    until    \  .•nii-i.-nhn 
syst..!.'    has   uiv.ii    pla.'.'    I..   .iia>t..|.',   lalimit    t».2   s mji,    tin-   jMirici.'    1 


ait.  ri.  -      ..,    (,t    III. 

Iil.   «.,ll  -.1  111,  1, 


""',  ",'    '"  '"   •',■"    '•    "•''■    l'>    l"-l'.">'   l""k-  ..Ml ,    .!..»„    ,!,.■    I.Tv.r   X 

',         ""    >'■"!. I.-   1. 1    ,,..,,.,„,,  ,,  ,,   ,,„|,    ,   ,,,,,|,   |,,,|    .|ir..i|>    tl 


TIIK    \(TH>\    <iK   TIIK    IIIMll 


14!) 


l|i<'n'ri>ri'   kIiIII    iiIV   rinlll    tlir    \  I'lltl'it-lc       Tin*   i'\ili-l    liloiui'iit    ill    iliitstnlr   iit 
itliirli    llli>   two  cilvilirH  lire   nuilill    lit'iillt;|it    iiiti)   riiiiiliimiii'iitiiili      it'.,    till- 

\ I'liti'ii-iilai'  MilvcH  iipt'ii  is  iiitlii-iiti-il  liv  till-  iMM'vc'i  I'liiiiiiiLr  luirctlKM'. 
llavinLT  tlills  ilrlcl'iiiilM'il  tlic  ixiict  IMoliii'liN  ii|'  )i|ii'ilili*_'  <llii|  cliiHililt 
!•{  till'  iiiirifiilovciitiii'iiliir  v^iUi'.  ««•  iiiiiv  imw  |mim'it<|  \„  i'ii|ii|ijii'i'  tin* 
iiitriivi'iitriiMiliir  ih'i'mhiu'c  nirvi'  w  itii  tliiit  tiikcii  I'roiit  the  iioi'tii.  Al'tcr  tlio 
iii'rrs>Miry  I'iililniitinii  iMiiii'd ions,  thin  cilixr  liiiN  lici'ii  |iliit'i'<l  ill  l-'iif,  ;t4 
HI  its  trill'  ifliitii'iiship  111  till'  vi'iiliiciiliif  cmvc.  Ui"_'iiiiiiiiif  ntfiiiii  nt  tlii' 
iiitl  (if  iliiisltili',  Ml'  liiid  flijit  llif  niirtii*  |>iTssiiri'  is  xciy  finiHili'iiitilv 
.'iliitvi-  tliat  of  till'  \  t'liti'iclfs,  iiiiliciilint;  lliiit  tlii>  M'liiihiiiiir  Milvrs  must 
I'l'  I'liiKcil:  mill  it   will  111'  t>I>st'rvul  iK.il   tlif  iiitriiM'iilririiljir  |iirssiiii'  iil 


I'lK  U  rus^iiir  vuivi-  .it:. I  luitui  ki.ftii.ili  .|  li.i\,  l.tvii  -tiin  1  irin-'-^i  .1.  'I  !u  |.rt  •.jili.'.titiiii . 
-I'lvuMiK  .iM'l  f"»t-|'li>Kl""  |-»i""N  'il  \i  iiti  h  illi'r  ^>-lMlr  .iu  "h'ivMi  li\  III.  v.itu.il  liiii  N.  "nu- 
ll    ».iln.   .I..M    .11    liir    l.r.t    iiiii.      Thr    ^i..ili.     \..;v    -    ..1.11    .11    til,     .,,,.,1.1    |i,i,    .,i,.l    ,1..<,.    aciiil    .it 

111.     Ihit.l    lin.        Til,'    .11'    ijKi«    ,.i.in    at    t'.i,     l..,nlli    !in,  .       Tin     |.i.-iti I    III,     t«i,    iiuiiii    liiMit 

i...|>    I-    .il...    iii.li,  .,i,,l. 

till'  liiLriiiiiiiii;  tif  svstnlc  tines  nut  lisf  siillififiillv  tn  ii|it'ii  tliiiii  until  an 
a|i|irti'iaii!t'  interval  !(l.(l"_'  In  (Ml4  seet.iiil  alter  liie  elnsurt'  nt"  the  aiirie- 
iilipveiitrieul.ir  vaKes;  lliat  is  In  sa\ ,  tliere  is  a  peiiinl  at  tlif  lienimiiu',' 
111'  vi'Mtrifiilar  svstnle  tluiiuf;  w  liieli  tlie  xenlliele  is  a  eltisetl  ea\  itv.  Il 
is  ,'i  I'fiititl  tliiriiii;  wliieli  the  \  t'lit  liele  liy  its  emit  laet  imi  is  '.'eltiiij.'  up  a 
^uriifieiit  aiiiiiiint  tif  pressiii'e  ill  the  lluiil  etintaiiiitl  in  il  Iti  I'tiree  open 
'lie  sfiiiiliinar  \;il\es  a<_'aiiist  the  resistaiiee  tif  the  pressure  in  the  Jitu'ta. 
:nii|  it  has  heen  ptipiil.ii'iy  eailetl  "the  iieritnl  tif  tretlinu'  up  steam."  or, 
in  |iliysiiilii<:ie  lan^ruatri'.  the  isdimtric.  tir  the  /in  siiln/i/niic,  perititl.  We 
■■liall  Use  the  last-iiieiititiiM'ti  teriii   in   tiiir   t'urther  tliseussimi   here. 


I    f| 


oiuJt^M^^ 


MICtOCOPY    RESOIUTION    TEST   CHART 

ANSI  ond  ISO  TEST  CHART  No    7 


1.0 


I.I 


1^:121 


2.2 


I:  1^     — 
::  lis  IIIIIM 


11.25   mil  1.4 


1.8 


1.6 


^     APPLIED  INA4GE 


!;■)() 


Tin;    liHCI  I.ATKIN    OP    'llli:    lil.ddl) 


'4 


Alter  the  inn'tic  Mihcs  liiiM'  liccii  oiH'iicd.  it  will  l)(>  <il(,>»efveil  tliat  the 
.(■sMirc  ill  tlic  vciitiiclfs  is  jiist  a  little  aliove  tliat  in  tlic  aortii,  and  that 
1  conlinnes  so  <l\irin<r  tlic  wliole  ot'  \('ntri(Milai-  svstnlc.  When  diastole 
sets  in,  tlie  |)i-essni-e  in  llie  ventrieles  (|uiei\l\  falls,  and  a  point  is  soon 
I'oaeliod  at  wliieli  e(|nality  ot'  pressure  in  ventriele  and  a<n'ta  is  a^ain 
attained.  This  eoiresjionds  to  the  nionu'nt  if  the  closure  of  the  semi- 
lunar valves.  The  pressure  in  the  ventriele,  althousih  now  rapidly  fall- 
in;;,  takes  a  little  time  before  it  has  fallen  low  enoutrh  to  permit  tlie 
auricular  valves  to  ojien.  Here  a^'ain.  then,  the  ventriele  is  a  clo.sed  cavit.v. 
and  we  have  what  is  knou  n  as  tin  ijuslsiilnii/niir  jxriud. 


CHATTKR  XVm 
THE  I'lMIMNti  ACTION  Ol"  TIIK  IlKAUT  (CoiitVh 


THE  CONTOUR  OF  THE  INTRACARDIAC  CURVES 

The  Ventricular  Ciirve 

Fnmi  ill)  aiialvsis  i>t'  tlic  (Miiitoiir  (if  cacli  cuivc.  t'liftlicf  interest iiiir 
l)(iiiits  are  broufrht  to  lijiht.  The  ventrieular  eurve  in  tlie  diairrani  alluded 
til  al)<)ve  (FifT-  •f4i  is  sliowii  as  liavins;  a  Hat  top  or  i)lateau.  ISy  the  use 
ot"  tlie  more  modern,  optieally  I'ceordiii^.  instiunients  it  has  heen  shown 
that  tliis  plateau  lieeoines  displaeed  liy  a  p<'ak  if  evei-y  preeaution  is 
'akcn  to  prevent  (liillinfr  dow  n  of  tlie  pressure  chan-res  in  the  instrument, 
;is  hy  opening;  wide  the  stopeoek  in  the  instrument  (  Fi^'.  :>;t).  The  peak 
IN,  liowever.  I»y  no  means  a  sliai'p  oiu",  so  that  we  nnl.^  fitly  deserilie  the 
I'liiitour  of  the  ventrieular-  cnrxc  durinir  the  s|)hy«;nii('  period  as  emisist- 
intr  of  a  risiiif/  porliou,  almost  continuous  with  the  eurve  duriiisj  the  ))re- 
spliyjiuiie  period,  a  .sinimiil  ami  then  a  ili rlliiiiii/  iinrfliDi.  which  is  usually 
slower  than  the  aseendinv'.  The  practical  value  ai'isinjr  from  a  study  (d'  the 
curves  lies  in  tlie  insifiht  wliieh  they  u:ive  us  into  the  nature  of  the  stroke 
nf  the  {-ardiae  |)ump,  The\  show  us  tlwil  the  impulse  \\hi(di  tlu-  ventricle 
Liivcs  to  the  moviujj:  mass  of  Mood  in  the  aorta  is  rather  a  sudden  tlmn  a 
^ustained  one.  The  column  of  liiood  in  the  aorta  is  a  miirhty  thiuf;  to 
move,  ami  it  xvo\dd  apjiear  as  if  a  sustained  pressure  l)rou>.'ht  to  hear  on 
it  durinii-  the  sphytimie  iieriod  wonld  he  far  nmre  etiieient  in  hriniriui: 
aliout  an  aderpuite  movement  of  the  hlood  than  a  sudden  jerk.  In  elosinjr 
a  heavy  <iii\v  a  slow  sustained  pressure  is  far  more  elTective  than  a  short 

{Misil. 

It  is  further  <d'  interest  to  note  on  I  lie  intraventricular  pressure  eurve 
tiial  tlier*'  is  very  little  indication  of  any  stioiKhirii  irans  iir  rihnitioii^ 
iit  the  moment  duiiiis:  which  the  semilunar  valves  are  opened  or  (dosed. 
.Nevertheless,  hy  dose  scrutiny  it  can  usually  he  seen  that  a  sliLrht 
'liiinije  in  the  direction  of  the  aseendiny  curve  is  evident  'vhen  the  \ah  .'s 
iipcn  (see  Kijr.  :i:{),  and  similarly  that  the  moiiiiiit  of  closing'  is  indicaJcd 
liy  a  sharper  lieiid  in  the  eiiive.  As  a  matter  of  I'act,  \Vii."_M'is  has  shown 
lliat  the  exact  contour  of  the  eurve  durinir  the  spliv  siiiie  period  depends 
ji.'irtly  on  the  decree  "f  scnsitixeness  of  the  uptical  manometer  i|se<l  iiiid 
linrtly  on  the  tension  e\istiny  in   the   veiitriide  just   hefore  contraction. 

151 


ui 


i:>L' 


Tin:  ciKci  i.ATioN  (IP  Tin;  iii.odi) 


In  tlic  ciisi'  (if  the  ri-ilit  vciitricli'  tlic  cdiitoiir  of  the  ciirvc  iiIsk  dcpt'iiilN 
(III  the  ilfjji'ci'  (if  li'sistillicc  to  llif  liliH'dlliiw  tllldlljrll  the  lilllllliili;if> 
ciicilit.  Till'  t(i|i  III'  llic  cili'vc  lii'ciHiH's  liiiijidci'  wlicii  the  iliiti.il  tiiisiini 
is   lii;.')!.   iiiiil    miiic    i'ihiikIi'iI    wIkm   tlicrc    is  :i    lii<rli    |iuliii(iii!ii'v    rcsist.iiii'i'. 

Aiiiiliicr  imiiit  111'  interest  in  (•(nincetinn  \vitli  tlie  ventriciil.ir  curve  is 
tlijit  cii'ly  //(  iliiishil,  II  ills,  I  nils  liihiir  llii  liiii  iif  :<  ri)  i>rissuri,  iiKlicitiiii; 
tli.'it  11  iiejiiitive  (If  suction  picssurc  must  exist  in  llie  ventricle  at  tiii^ 
time.  It  will  lie  liiither  (iliseived,  Iniwever,  tliiit  tiiis  siil)atni(is])lieii(' 
pfessiiic  exists  I'm-  (inly  a  very  sliiirt  time.  Tlic  aiii-iciilnvcntrlciilar 
vahi's  IiciiMj-  (ipeiied.  a  similar  iiciralixc  iiressnre  is  also  ])i'esent  in  tlie 
aurienlar  traeimr.  Were  we  to  depend  mi  such  rccoi'ds  alone  fur  evidence 
of  tlie  actual  existence  of  tliis  iicirali\i'  pressure  in  the  heart,  olijectioii 
miirht  lie  taken  to  the  coiKdiisioii  on  the  urmind  that  it  was  due  to  ine 


to  rr^^ncrr.H^r 


irjx  \a.je 


Viil.     ,!.',       \'.in     l-i.iiik'-     iii...iii.,il 
liil.i-     I'lUviiii     In  .lit    :i:iii    lllil<;iry     n, 
ni.iMnMiiM.    niitiininip,   m    <.ii!M!ai>    mm 


to  hearl 


1..1     \.|K..     ttlinli     i-     |i|,i>i'l     ill     Ih.-     (..iirM-     1.1     !!■, 
!'.>■    tin  mm;    iIk     si..i.,  i.,k>,    it    iii.iy    In-    llMil    ii-.    . 
inl-.i!    tnln>    ..1.(11'.      ll-'i'Mi;    StiitluiK  i 


sudden  I'ccciil  til  which  the  iiistnimeiit  is  sulijeetcd  at  the  licfjinninsr  ol 
diastole.  It  is  necessary  thcrcfure  to  control  these  (iliservalions  liy  thi 
use  of  an  entirely  dilTereiit  method.  This  consists  in  conncctiiit;  tin 
heart  with  a  ralnil  nnrrnrii  iiiniiiiiiit  li  r  i  see  l-'i;;.  :!.")'.  This  iiistru 
meiit  does  not  of  conise  record  any  sudden  chanyres  of  pressure  in  th' 
cardiac  cavity.  Inif  in  oliediciice  to  chaiifres  in  ]iressnre  the  luereiiry  sl(i\vl> 
iiiiives  in  the  (lirccliim  in  which  the  valve  permits  it  to  move.  Such  ai 
instrimiciit.  with  the  valve  ojieninij-  towards  the  hear!,  is  called  a  iiiiliiliia: 
iiiaiioiiietir.  and  after  it  has  licen  connected  with  the  ventricle,  it  will  1' 
found  that  a  ncyiiti\('  iiicssure  of  perhaps  4(1  or  (in  nim.  11^  is  recorded 
Evidently,  then,  tin  ii(;/iilici  iirissiirf  iluis  itrlioilhi  i  rist  in  tlif  r(  nlrid 
dnrinjr  some  phase  of  tl;e  cycle,  and  the  (piestion  arises  as  to  whether  r 
i<  of  inipiirtaiice  Isi  cmincctiiiii  with  ilie  ptiiiipin;:,'  acfion  of  tlic  heart,  ,\' 
first   si^hl,   considcrin;.''  the   heart   as   an    elastic   slriielnre,   we   mi^rlit   coi! 


Tin:  I'l.Mi'iNci  Afiiox  OK  Tin:  iii:.\i{i 


!.-);{ 


tli.it  tlic  licir;itivc  incssurc  wciuld  scivc  to  siii-U  lildiid  into  the  liciirt. 


.jiNt  iis  It  Slicks  water  in  nii  nnlinarv  luill  svrini'c.     C 


iKscr  roiisM 


Icriiti 


ill,  liciwcvcr,  siiow  that  this  (•(.iicliisioii   is  iiiiti'iiai)Ic,   partly  hccaiisc  tin 


iH'LMtivf  in'fssiirc  exists  ill  the  veiitriele  I'cir  sti  short 


le  veiiirieie  lor  so  siiort  a  period  ol  tune,  ami 
IMTtly  hecaiise  it  would  have  to  operate  on  th"  slowly  moving'  eohiiiiii  oi' 
Mood  in  till'  thill-walled  veins,  with  the  result  that  it  would  eaiise  ih(.  walls 
of  these  vessels  to  eoiiie  to;:etl.er  lather  than  produce  a  iiioveiiieiit  of"  the 
IpIoimI  eoiitaiiied  in  theiii.  The  iieuative  luessiiie  of  the  heart  can  not 
llierefoi-  lie  of  iiiiich  coiiseipience  in  atlraetin-.'  the  venous  lilood  into  the 
Vfiitricle. 

Several    factors   may   cooperate    to     produce     this     iieu'.itive     inessiire, 
.iiiioM!.'  them  heiii^'  the  sudd. mi  o|)eiiiinr  out   of  tiie  hase  of  tlie  ventrieles 

.It  the  heirinnintr  of  diastole,  the  elastic  r ii  of  |h<>  tiss.ie  which  hecoines 

compressed  in  the  heart  walls  diiriiin'  sxstole.  fh,.  tury:eseeni'e  ai'  the  walls 
Hi'  ihe  ventricles  produced  hy  the  sudden  iniiish  of  hlood  into  the  coro- 
nary vessels  at  the  li  iinniii^  of  diastole,  all  of  which  factors  tend  to 
ciiise  an  ojieiiin^'  out  of  the  \valls  of  the  ventricles  with  a  coilse(ilient 
increase  in  the  capacity  of  llieir  cavities. 


The  Auricular  Curve 

i;\amiii;itioii  of  the  iutraaiiriciilar  iiressiire  curve  is  of  i>articular  iii- 
i.rrsl  I.ecaiise  of  the  rclal ioiiship  wiiicli  it  has  1..  a  traciiifr  taken  of  the 
'iiovements  in  tlie  .juj-uliir  vein  at  the  root  of  the  neck  (see  ])ajri'  274>. 
Tliis  .jiiirular  pulse  curve,  as  it  is  called,  is  luoduced  mainly  l>y  the 
ili,iiii.'es  of  pressure  occiirrim:  in  flie  iinride.  from  which  it  <lifT.'rs  only  in 
I  lie  relative  lieii;ht  of  the  various  waves.  ])y  <rradua1in!r  the  iiitra- 
■luiiciilar  i.ressiire  curve  i,y  ihe  method  deserihcd  ahove,  we  can  tell 
•  AMctly  the  mauiiitiide  in  the  cliaiiye.;  ,,f  pressure  occiirrin-;  diirini;  oacli 
c:ii(|;;ic  cycle.  TIlis  obviously  can  piit  he  done  w  ith  a  tracing;  taken  from 
'III'  .juiriilar  vein,  ahhoii-h  (pialitativdy  the  tracin-;s  relh.ct  exactly  the 
'■'iMiiv'cs  that  are  oceiirrin>r  in  the  auricle. 

•hi  examining'  the  auricular  iiressiire  curve  (consult  Fi(rs.  :U  and  it"),  we 
liii'l  that  after  the  wave  of  |)resystole.  which  of  course  coincides  exactly 
"Ith  ihat  on  the  intraventricular  .  irve.  a  second  wave  occurs  .•nlminatiii'.' 
Ill  ii  peak  almost  exactly  at  the  lieiriiiiiinii-  of  tli.'  sohymnic  jieriod.     The 

■  irvc  then  raiudly  descends,  usually  ind 1  he!ow  tlie  lii f  zero  i.res- 

~'iie,  and  slowly  rises  throuudioiit  the  rest   of  ventricular  systole,   until 

*'"■  ' '"'"'  "f<'I)fiiiii!,'of  the  aiiriculoveiitricular  valve,  wlien  it  d.-'scends 

.I'-'iiiii  and  thereafter  runs  paralhd  with  the  ventricular  curve.     The  let- 
'"!'•>  used   to  desimiatc   the   waves  are   the   same   as    ihose   eiii|iloyed    for 

■  iiiiliii'   waves  shown   on   the   .jufrular   pulse   tracinj;,   and    altli.nii.di    the 


M^Jt^'* 


U    i 


1.".4 


Tin;  ciHii  i.AritiN  ok  tiii:  iti.(M)i> 


^     I 


IcttciiiiK  i^  more  m-  less  jirliit  iiii'\ .  wv  imist  iicccpt  it  lifcause  of  its  nfo 
criil  nxiMii'  in  all  \\«vk  «(  tliis  Uiiul. 

As  to  tlic  ciiuscs  ot'  till-  waves  .1  is  (if  i-mirso  caused  hy  amicular  svstui 
«.r  piTsystdU-:  ('.  ((ccuniMjr  as  it  ijocs  at  the  lii'};iiiniiitr  of  the  poriotl  > 
vfiitiii-ular  sy>t(il<',  is  caused  liy  tlie  l(id!.'iii;;  into  the  auricle  of  the  elosi .; 
ainicnioventiicular  valve.     Tiie  lloor  of  tlie  aiiiiele.   in  otlicr  uoi'ds,  ii' 

this  1 t  lieeoiiies  soiiiew  hat  elevated  and  iiiiimtts  to  the  blood  whid 

is  lestiii!,'  npon  it  a  slii;ht  wave  of  pi-essiii-e,  \vhicl>  is  transmitted  aloirj 
the  veins  t'or  a  eonsideralde  distance.  The  siicceedinfr  dei)ressi(iH  i- 
marked  .'•.  and  the  iiejiative  pressure  u  hicii  it  indicates  is  ])rohahly  dii' 
to  the  CO  operation  of  three  forces,  all  tendinj.'  to  increase  the  a»irie>d:t 
capacity:  iP  the  diastole  of  the  wails  of  the  auricle:  (2)  tlio  desceii! 
of  the  auriculovenlricular  <;roove,  thus  tendiu}:  to  open  out  somewhat 
the  folds  in  the  walls  of  the  auricle:  and  C^i.  i.rohahly  most  important  <•' 
all.  the  tendency  of  the  thin  walled  auricles  to  liecome  dilated  as  a  resul' 
of  the  sudden  diminution  in  intrathoracic  pressure  produced  at  each  hear' 

heat  liy  thi'  discharire  of  lilood  from  tlie  heart   and  intrathoracic  hi i 

vessels  into  those  of  the  rest  of  the  hody.  All  thin-walled  structur.. 
in  the  thoracic  cavity,  the  auricles  iuehuled,  will  expand  to  take  up  tin 
cMra  room  created  in  the  thoracic  cavity.  Similar  nejiative  heart  juilsc- 
as  they  are  called,  can  lie  observed  with  each  systole  in  the  luufjs  ainl 
in  the  es<iphM'-rus. 


THE  MECHANISM  OF  OPENING  AlfD  CLOSING  OF  THE  VALVES 

When  physical  \.ilves  open  and  close  as  a  result  of  the  changes  ill  pn 
sure  on  their  t«o  surfaces,  a  certain  amount   of  lluid  must   succeed 
l)assinff  the  val\e  Haps  before  these  become  iierfectlx    closed.     But  tin    ' 
is  every  reason  to  believe  that  such  is  not  the  case  in  the  heart,  the  flai'- 
of  both  the  auiicidovcntricidar  and  the  semilunar  valves  beins;  alrea^l 
completely  closed  before  pi-essurc  conditions  entiiiliuji  a  jiossiblo  rejin 
;2itation  of  blooil  thiouuh  them  become  establislied. 


Auriculoventricular  Valves 

Durint:  diastole  the  flaps  of  the  auriculov  .■utricular  valves  are  hanu  - 

dovvii  into  the  \cntriclc  and  lloatinv'  in  a  half-o])en  position  in  the  bh'  i. 
which  is  meanwhile  .•iccumnlatinsr  in  the  chamber.     This  position  is 

pendent   upon   the  opci;ition   of  t\\o  opposinj.'  foi'ccs  on   the   valve  tb!;  -: 

the  jiressurc  of  tlie  blood  lloHinf  from  the  .luricle  oh  their  upper  aspc  ~ 

and  rctlccteil  wa\es  of  pressure  from  the  walls  of  the  ventricle  on  tl  ' 

under  aspects   i  centlipetal    relluxi.      When    presystole   occurs,   the   p  - 

sure  of  tile  auricular  stream    momentarily    iinMcases,   thus  slifihtly  •  - 

teniliii''  the  vvall  of  the  meanwhile  relaxed  ventricle  and  after  a  moiiifi  > 


THE   I-1MI'IN(1    A<TION    OK   Till;    IIKAKT 


155 


.|i  liiy  cnusiiijt  the  icflpctod  wave  to  lircomc  more  |)ioiiouiict'(l.  At  the 
■iame  time  the  nuisciilar  (ihers  in  tlie  valve  Haps  ( Kiirsehnei-'s  filters) 
iMiitract  and  make  the  (laps  shorter,  the  total  etTeet  of  the  two  faetois 
liiinir  that  the  valve  takes  up  a  position  iieaier  that  of  elosure.  When 
piTsystole  suddenly  stops,  the  letlexed  waves  will  jiersist  for  an  instant 
(if  time  longer  than  the  aurieular  wave  whieh  eauscs  Iheiu.  Iiecause  of 
I  he  elastic  nature  of  the  veiitrieular  wall,  so  that  the  valve  flaps  close 
with  perfect  opposition  not  merely  at  their  edfjes  hut  also  for  a  con- 
vidcialile  distance  aloiif;  their  ii]>per  surfaces. 

When  ventricular  systole  starts,  'lie  only  cfTect  of  the  hi;rh  pressure 
\\  liich  is  brought  suddenly  to  bear  m  the  under  surfaces  of  the  already 
iloscd  valves  is  to  cause  them  to  vibiMte  and  to  bulfte  into  the  auricles, 
ticinfj  meanwhile  anchored  down  and  i>ii'vented  from  flapping  into  the 
auricle  by  the  chonbe  tendinea".  Althoujfh  there  is  reason  to  believe  that 
the  musculi  papillares  to  whieh  these  are  attached  besrin  to  contract  at  the 


he    .Vi. — Diagram    to   ^how   the    fnsitiotis    of    the   cardiac    valves:      1.    diirinR    diastole;    J,    duniiK 
the    presphyginic    period;    .'.    duriiiR    the    sphygiiiic    period. 

NfTv  outset  of  ventricular  systole — indeed  slightly  to  precede  it  (see 
iMtrc  2W-i).  ami  thus  keep  the  chonhe  tnut.  yet  as  systole  continues  the 
iiiiitraction  of  these  muscles  becomes  nnu'c  and  nioi'c  pronounced,  and  the 
rosiilting  tightening  of  the  chorda-  serves  to  draw  (b)Wii  the  valve  Ibips, 
so  that  progressively  larger  proportions  of  their  upper  aspects  tend  to 
liccome  opposed.  Meanwhile  the  anriculoventricular  orifice  is  also  be- 
'oniing  narrowed  down  on  account  of  the  contraction  tif  the  musculature 
"f  the  auriculoventricular  groove. 


I  t 


Semilunar  Valves 

The  mechanism  involved  in  the  operation  of  the  semilunar  valves  is 

■inewhat  different.     It  has  been  shown  that,  when  fluid  is  flowing  in  a 

iiic,  the  pressure  and  velocity  are  not  equal  in  the  axial  and  peripheral 

'  Its  of  the  stream.    In  the  axis  the  velocity  is  greater  than  in  the  layers 

'    fluid  next  to  the  walls,  but  the  pressure  is  less.    These  facts  can  be 


I     1 


l.-.d 


Tin:    rlUCI  I.ATIIIN    (IK   Till:    l'l.(H)l> 


(li'iiiiiiisliiilcil    li.\    iilisciv  iiii;  till'   (low    |l|iiiut:li   M    uiili'   lillic   III'   WMtcr   i' 
wllicli    .-lie   sii^pi'iHlcil    lMMi|m(lilllil    s|iiiirs.      liy    ])liH'illi,'    ill    llic    llllii'   siii.i 

Im'iiI   IuIms  >ci  iiii.um.il  tlijit  "111'  iipfii  mil  lirs  iii'iii-  the  iicriplicry  Miii 

llii'  niliir  in;ir  lli iitiT.  it  ciiii  l)i'  si'i'ii  lliiit  llic  (lilTi'iciirfs  in  prcNMH' 

iiri'  siii'li  iis  til  I'iiUM'  till'  lliiiil  to  \\«\\    Irniii  iH'iiiiiiciy  tn  iixis  (ci'iitriprt;  i 
ii|ilii'>  I. 

ir  till'  lii'iit  tiilii-.  iiiT  ii'Mil  til  stmly  till'  iiiiiililiiiiis  111'  How  ill  ii  tiiln'  wliiri, 
Miililnilv  In iiir>  w  iiliT,  il    \\ill   111'   fiiiiiiil  tiiiit    wlii'i-i'  till-  wiili'  iiiirti.iii 

st.■ll■t^   iililliprlill    iililics   .irt'    set    lip,    wllicll    tcllil    tii   I'.'IITX     tllC   si'Cils    ilit  ■ 
tlir  :i\is  111'  till'  >1ii';iiii.   w  lii'i'i'  tlii'ir  vi'lnrity   is  irrciilly  iiii'ii'Msi'ij.      .\n\ 
llicM'  .-111'  till'  i-iiiiiiitiiiiis  iilil;iiniii'_'  iit   the  licuiiiniii^  of  tlu'  liir'_'('  .'iitciir  ■ 


S.a.-D.v 


I  l|.it;i  .i-'i     ^ti  .u  MIL'     til'      I'.^il !     ii 

I  S  .1        1 1.\  J     anil     il'iii  iiii.'     tin- 


111'    ihr    lli'.irt,    tl lilii-r    ililii    tlir    \  I'llt  rii'lcs    lii'ill'/    ciilistrii'tril,    wlliir 

till'  >ilills  \;il>:il\;i'  lln'  vcsmIs  ;iri'  dililtcil.  A  crlit  lipi'Iill  Mirti'N  IllIK' 
M't  u|i  ill  till'  siiiiiN.  ti'iiiliiiii-  til  tliriiw  till'  \;il\i'  lliips  iiitu  :i  cIum'iI  }>' 
tiiiii.  wliii'li.  liiiw  i'\  (I-  is  pr('\  I'liti'il  liy  the  liliiml  nisliiiiu'  lictuii'ii  tl 
I'liiiii    till'   \  clitrii-li's.      'I'lii'y    thus   tiiko    up   ;i    liiiil-pnsiliipii    ;iiii|    \ilii;ili 

till'    slllillll.       Wlllil    tllC    I'lllUN     tVnlll    the    VCIltl'il'll'    stops    lit     tilt'    ('Mil    111'    - 

toll',  till'  irlluN,  Instill'^  I'm-  il  iiionii'iit  loii'ji'i'  Mini  lii'iiiir  now  iiiiopp"- 
iiiiiii('ili;iti'l.\'  rlosi's  till'  Milvi's.  ill  wllicll  position  tiicy  urc  thru  iiiiiintiii' 
liy  till'  <rii';iti'r  pri'ssiiii'  on  tlirir  iippiT  siii  liifcs. 

Tlir  position  of  till'  \;il\('s  icl:iti\i'  to  tlii'  I'M'iits  of  tlic  I'iiriliiir  i-yi-l 
sliow  n  in  l'"ius.  :l(i  niul  -iT. 


Tin:    ri   Ml'IN(i    .\<  TION    OF   Till;    IIIAUT 


THE  HEART  SOUNDS 


157 


Diiiiiisr  <'i'rtaiii  i>li!iscs  of  tlio  cyclo  distinct  sounds,  tin-  heart  sounds, 
i'.in  lie  hcnid  liy  !i|ipl.vint;  a  stetlioscoiH.  to  tlio  llioiacic  Mall.     Tin-  first 

iii>>  .'it  tlic  liciriniiiii};  of  vcntricnlar  systole  and  is  liest  lieard  ()V<'r  the 

,i|ii\  licat;  the  second  occurs  at  the  lpc<;innin;,'  of  diastole  and  is  lieard 
l.^t  al  the  second  riu'lit  costal  cartilage  or  in  the  second  left  intercostal 
-[lacc.     A  third  sound,  nnidi  less  ilistinet,  is  sometimes  heard  in  dia.stole 

,1  sli(,it  time  after  the  s( nd.     To  study  the  exact  time  relationship  of 

'li''  >' 1><  the  vihialions  which  they  set  nj)  can  he  re<'ordcd  irraphically 

iilMUiiside  cardiac  tracinu's  hy  means  ,i(  a  microphoMe  attachment  to  the 
i-li'i-l  roiMrdiojfraph   i  see  p.i^e  'J.'iil). 


Causes  of  Sounds 

It  lias  lieen  found  that  the  jirsl  sukihI  consists  of  two  distinct  elements, 
I'rii'  hiuli  pitched  and  the  other  of  a  dull  ciuiractci\  The  fornu'r  element 
i-  liclicved  to  he  the  result  of  \  ilir.it ions  set  up  in  the  Haps  of  the  auiic- 
iilnMiitricnlar  valves,  and  theichoe  in  the  Mood  in  tile  heart,  hy  the 
sii.ldcii  rise  in  systolic  pressure.  The  dull  element  on  the  other  hand 
K  iiiiiloiihtedly  of  muscular  oii'.'iii.  The  evidence  for  these  conclusions  is 
ti-  I'nljows:  (li  When  the  auriculoventricular  valves  are  )irev(nted  from 
'I'.siii^  pniijcrly  either  hy  disease  or  liy  ])usliiiiir  a  loo])  of  wire  down  the 
l.ii'.'c  veins,  the  hi^li  pitched  (piality  disai)i)ears,  and  nothiiiy;  hut  a  rush- 

'   -  ^"iind  ;i( mpanies  the  dull  limit  produced  hy  the  coiitraclintr  muscle. 

-  In  a  heart  that  has  lieen  rendered  l>h>o<lless  hy  an  incision  iteai-  the 
■  '{■■■\,  or  even  in  an  excised  Init  still  h.'atinir  heart,  the  dull  eleiiient  id' 
!l  '■  liist  sound  still  eoiitinucs  to  l)e  heard  f,,r  a  short  time.  Tiiat  eoii- 
I  •I'-'iiii;  muscle  prodni-cs  a  sound  is  a  wcll-estahlished  fact. 

'Ilicic   arc.    however,    many    oli.<euie    phenomena    connected    with    the 

'■■i  i-.iliiiii  of  the  liist  sound,  lint   we  can  not   u'o  into  such  controx  eisial 

•..Mils    heie.      .\    close    inspection    of    the    elect roiihonojiiaphic    tiacin",' 

-'   '1'.  -  that  till-  sound  starts  at  the  ln'^-inninir  ,,|-  il,,.  |)iesphys;iiiic  period, 

■il   tli^il    it   lasts  with   madiially  decliniiiir  variation   in    intensity   until 

•  !i  iiilii  ih,    sphynniic  period   i  l-'iir.  :!Si. 

""■  ■"•    ""'   <■■"(/</  nis  ■• iirately  at   the  heirinninu'  of  diastole  .■ind 

'I  '•■;,, lily  he  shown  to  he  caused  hy  tli.<  sn.jden  vjiuttinu:  and  stret<'liin!,' 
'lie  s.iniliinar  vahes,  which  throws  them,  the  hlood  in  conlaet  with 

'■'    •'""'  '• eiifhhoiinir  walls  of  the  aorta   into  vihration.     Proof  of 

Insioii  is  furnished  l.y  the  following'  facts;     The  se.'ond  sound 

li'if'l.v  disjippears  if  the  hlood  i^  let  oil!   of  the  he.-irt  hy  oiieniiiir 

•i|"N.  iind  it   is  replaced  hy  a  nishilii:  "limit"  if  the  llajis  are  |)re- 
•■■'I  fioiii  clnsin^r  as  a  result  of  di.sease  or  of  hookinj;'  them  hacl<  hv 


II 


I.  i 
I 


F 


I'lS 


TIM     run  I  l,\TU>\    OK    llll     Itl.iMili 


jmHsiiiL'  a  will'  ilowii  llic  ciirolid  iirlcn.    Tlif  Ihinl  ^oiniil,  alllioiiuli  nii.l 
lilc  only  ill  stnnc  iiulix  kIiuiIh,  mil  ni'vortlii-h-ss  lie  shown  to  cxImI  l>y  tli. 
(•I.Tfrn|.ln.iio^iii|ili.   anil   siii.c   it   or.Mirs   at    tin-   tinu'   when   tlic   anrifiil. 
\fntiitiilai   valves  oiicn,  it  is  liflitvi-.l  to  ilf|icnil  upon  tlif  smltU'n  iiiiuvl 
of  Idooil  Ironi  auiirlcs  to  vcntiififs. 

The  Ki«'atfst  iiiiport.iiu I"  tiif  sounds  is  in  tin'  clinical  diairiioHis  of  \ai 

vnlar  ami  other  lesions  of  the  heart.  When  a  valve  leaks,  for  exaiui)!' 
the  l.loo.l  escapes  past  it  under  jfieat  pressiiie.  and  is  ojoeted  into  n  iiias> 
of  Mood  at  low  pressure,  these  heins.'  conditions  wliieh  are  well  kiioui 
to  create  sounds  or  hniits.  \\\  examining  the  exact  relationship  of  such 
limits  to  the  normal  heart  sounds,  deductions  can  lie  drawn  concernin', 
the  condition  of  the  various  valves. 


Record  of  Heart  Sotmds 

The  heart  sounds  have  lieen  jrraphieally  recorded  liy  tl■anslnittin^'  thim 
Ihroutrh  a  stethoscope  to  a   microphone  placed   in  circuit   with   a   strin- 
■.Mlvanoineter  )  elect  rophonoirrains).      Through    this    circuit    jiasses    a    cm 
rent    the   strength    of   which   deiiends   on   the    resistance   offered   liy    tl.. 
microphone,  this  resistance  liein«  proportional  to  the  nunilior  and  ampli 
tuile  of  the  vihrations  of  the  sounds  transmitted  to  it  through  the  stetli- 
scope.    Tliero  are  several  objections  to  this  method.     One  of  these  is  d  ■ 
pendent  on  the  varyint;  distance  of  the  heart  from  the  chest  wall,  whirli 
causes  many  of  the  sound   vihrations  to  lie   lost    liefore  they   reach   tin' 
stethoscop.-;   another,   on    adventitious  sounds   arisinj,'   from   contraetiiiL' 
muscles,  the  iiniiacl   <if  the  heart   asrainst  the  chest   wall,  etc.,  and  sli'l 
another  on  uiie(|ual  resonation  liy  the  air  in  the  neisrhhoriiiK  portions  .1 
lunirs.     To  investisrate  the  prolilem   more  thoroughly.   WisiRers,"  usii- 
anesthetized  animaK,  has  recorded  the  sounds  liy  carefully  stitchiiisr  i" 
the  heart  (exposed  throuiih  a  small  opeiiin'r  in  the  pericardium)  a  levi  r 
the    end    of   which    was    attached    to    a    "transmitter"    consistinji    ot    .1 
small  capsule  covered  with  ruliher  dam.     The  transmitter  was  conned.  1 
hy  ruliher  tnhinf:  to  a  •recorder""  consistiiifr  of  another  small  capsule  can 
intr  on  it^  memhrane  (made  of  ruliher  cement)  an  eccentrically  placed  sin  M 
mirror,  »n  to  which  a  heam  of  li^dit  was  thrown.     The  movements  of  r  <■ 
beam  of  lijfht  rellected  from  the  mirror,  and  caused  by  the  sound  vil"  i- 
tions,  were  photoffiaphed.     Mechanical  vibrations  set  up  in  the  appar.i'  > 
itself  were  lar>fely  eliminated  by  a  side  oiieninjr  on  the  recorder,  and  ■    '■ 
elTect   of  outside   sounds  mininii/c.l   by   surrounding   the   recorder   b;    ;i 
ventilated  }rlass  housiii}.'. 

Althoujili  tliis  apparatus  is  not  free  ffoin  faults  due  to  iidiereiu  vi!  : 
tion  fre(|uency  and  resonance,  the  records  seemed  by  it  are  valuable  11 
showintr  the  exact  relationship  of  the  sounds  to  the  events  of  the  carci   «• 


Illr    I'IMMSII     \l  Tlli\    til-'     Till      III   MM' 


I'l'.t 


\rlr.  Till'  \  ilii'iitioii^  I'i'diii  till'  Iwii  Mill  fides  ni'c  aliki'.  Imt  ililTci-  fmiii 
icisi-  tak«'ii  from  flic  imrta.  'I  hi  first  n  iitrii  iilur  xniiinl  i'(iii>i>t>  iM'  fniiii 
!.■  to  tliiiti'fii  irri't'iilar  \  ilinititms.  iisiiallv  in  liner  u'i'i>ii|is,  the  lii'si 
>iii|iiist'<i  III'  two  sitiiill  \  iliriititiiis,  lilt'  iiiiililli'  mil'  n|'  si'mtji)  |jir)ri>  \  ilirn- 
niis,   iiiiil    till'   third    lit'   .1    \iii'\iiiu    iiuiiiIkt    <>(    mimI!    \  iln'iitimis.      'I'lir 


^l 


••   ,V*i  '.'V*' 


•AV 


I       I  II  II 

.  >  ~'«  f"  ^•' 


A 


'    A 


Yl^H^fPPH.    )|)^^^MM: 


I  IK  .ix,  l.li.lH.pi.MuKiani.  aliiMK  uiili  ir.ti.,v.  iili  u  rli)  pi,. MM,  ,inv,-  ti..„,  ih,,,-  ,lil 
•I     ixr-Timnits.       Ill    .1    iln-    vi|,|„in„i,i    lurn-    sli..«.    tlie    |.i..Mir,.    lli,-    iiM.l.ll,-    ,>i,,'    llu-    ^,■|IM.U 

I.'-  riKhl  viHiul..  .i,,,!  ll„-  l„»,.riii...t  ,.n,-  lli,..f  ..I  il,,-  :.„na  /■  „„l„,i„s  tli,-  Ml.itiv,-  |.,..i 
'.   '.    '""',  '"'Y'-''  '*    ''"'     '"    '""I'ai'Ual    ,is,,llali,,.i,.       ,s.'    ni,li.-,it,,    tlif    s,i-,,ii,l    soiin,!.    .in,l 

•■    •■    an,     V    the    ccrvctf.l    tiuir    nlatiiii;,    .,i    ll.r    1,1,1    v,,,,,„U.       |„    /(.    ||„.    prcssiir.-    an, I    s.>iiii,i 

.■  ar,.  l,„.h  trum  tlir  kit  v.-iilri.U-  ll.-iut.  -.i,,,,-  :,.  i„  .(1.  I„  c,  ,h,.  a,,,-,;^.  a,„|  |.„l„„,i,a,v 
'    ■.II   ."inul,   :,r.-   shi.wii    (Irtltr,   s.inu-   as    111    .11.      1  |-r,ii,i    \\iB«,,-   an.l    Hiaii.l 

Elation  of  the  sound  is  from  (LO.")  to  O.hVi  seconds,  and  the  iieriodieity 
"Ml  (1.004  to  0.054  per  seeond.     When  comiiared  with  .'in  intraveiitrieii- 


J 


ItiO  Tin;  <  iiiri  i.A'riii\  III''  'I'lii:  iii.ih)I> 

liir  |iri'>Miri'  riii-\c.  Ilii'  initiiil  viliniiimis  ncM-iir  (101  m'i'iuiiI  prinr  f 

: *i :..   .  :i *: I.li...  4l...:i. ii.i^i    tlitn 


II  till'   li-' 


llir  |iri'>Miri'  rllI'M'.   Illi'   inillill   Vllirillliins  nriMir  imm    mthmh   |iriiir   Ml   Mil'   M 
in  pliv-slU'i'.  the  lllllill  \  ilinitiiilis  li';i('liiliu'  their  irirjllrsl    lllil|tlitl|i|('  licl'ii 

till-  spliyumic   pniiiil   ip(iiiM>.  ami  tlir  linil   \  ilniitiniis  nrciiriinif  iliirii 
tlii'i'iiil.v  piiif  iif  till'  spliyu'iiiif  pi'ii'iil  iiiiii  liii'ii'lni'i'  just  lii't'ini'  tln'  iimtir 

pi'i'NNiiic  liiis   iciM'litil   its  lii'lulif.     Till'   iiifiiii   vilpiiitimiH  liicrcfuic   i ur 

lllllill!.'  till'  ilrsi'i'iiiliiii;  lliiili  111'  111'-  I!  wiivi'  111'  till'  I'li'rtriii'iiriliii'.'fiini  ili«' 
iriiiiiiii'.'  tl.lM  si'i'iiiiil  lii'I'iirc  its  rniiipli'tinii  > .  tin-  siiiiill  pi-iliiiiiiiiiry  vilir.i 
lions  iM'i-iirriiiif  iliiiiii'_'  tlir  iisi'didini.'  limli.     Wlirii  liiki'ii  t'n  in  tin'  iinrt.i, 

•  lie  II Ill  III'  till'  liist  siiiiinl  is  siiini'wliiit  ililTi'ii'iit,  tlii'ii'  lirinu  iin  inili.i! 

\  ilii'iitiniis  iiiiil  tlic  iiniiii  iiiK's  liciii'.'  uj'  irri'Mii'i'  lit'i|iii'iir.\  iiiui  ii'iii'liiii-j 
tlii'ir  iiia\iiiiiiiii  railiiT  tluiii  tlnivc  taken  I'mii  the  ventricle.  The  siili 
si'i|iii'iit  viliratiiins  are  alsn  laruir.  espeeiallv  when  the  aortic  pressiiii' 
is  lii>{li  (  \-'\i'.  :it^i. 

The  reconl  of  the  siciunl  sntiml  at  the  vehtriile  is  iinieh  simpler  an. I 
ilsiialix  III'  li'»s  .iiiiplitnili'  than  the  lilst.  cnnsistinv'  of  two  to  six  vihratiun^ 
lastiii;:  (Ml1."i  to  (IO."ili  siTiinil.  'I'liex  I.eL'in  a  slmrt  time  after  the  veiilr'rM 
Iiir  pressure  1  eyiiis  to  fall.  appio\iiiiati'ly  at  the  iliemtie  nnteh  of  the  ami  :.■ 

curve,  lieinn  ( ipleteil   ill   fn  in  (i.lll."»  to  U.irJ.'i  s ml  after  the  hottnni 

iif  the  nnteh.     Their  relatiiinsiii|i  to  the  T  wave  is  varialile.     Taken  fnu!. 

the  amta,  the   rri'inil   of  the  s mil   suiiinl   shows   viliratiuiis  of  irreali  i' 

aiiiplitmle  ami  of  a  '.'leatrr  fiei|ui'iii-.v    than  that    from  the  v  eiitriele. 


I 


ni.M'TKIi   \l.\ 


TIIK  M  TUITION  or  TIIK  IIKAWT 


^ 
s 


THE  BLOOD  SUPPLY 

li I<l  liloodc.l  iiMiiiiJil.><.  siicli  lis  tlic  (nm,  til,'  liciMf  iiiiiscic  is  iKnnisIicil 

'  ^    ''1 1  soakinir  into  if   IVom  (lie  licmt  cliaiiiltfis.  wliicli  iiiili'nl  ilu  imt 

"'iiii  ili-liiiit)-  <'iivitii's  lis  ill  tlic  iiiiiiiiiiiiiliaii  liciiH.  Imf  exist  us  nn  intcr- 

' "K"'  "f  Miusciiiiir  lisNiii'.     Ill  tlic  li.Mfts  of  JiiL'lii'r  iiiiiiiiiils.  tlic  iiiiisi'ii 

'..inic  is  su|i|)iifil  liy  spcciiil  iirtciics  (llic  ctiniiuiiy  i,  iiltli<»ii(rli  i rtaiii 

I'UMiiiit  111'  liliiiiil  iiiav  still  pass  (liicctly  iVoiii  tlic  canliac  cavities  into 
■..'■  iiiiisciilatiirc  tlif<  iijjii  tlic  xciiis  of  Tliclicsiiis. 

Tlic  Illative  imp  Mtaiicc  of  the  val•iou^  hiaiiclics  of  tin r  nary  nitery 

I     iiiaiiitaiiiiiitf  an  ailcipiatc  niitiition  of  the  heart   lias  I ii  stiidieil  Ity 

'l-civiiiif  the  ctVect  of  1 Iiision  of  one  m-  more  of  them  (W.  T.  Porter".) 

"■  •liisioii  of  the  ciiciiiiitlc\   lirancli   of  the   left    coronary   artery   eaiisod 

.i"'st  of  the  licartlical   in  alioiit   M)  pci nt   of  cases,  the  arrest  )mi\K 

'-"•■illy  II iiianicd   liy   (iliiiilaiy  contraction,     (tcclnsion   of  the   ri»{ht 

■■"!"Maiy  arrested  the  ventricular  coiitiiietion  in  alioiit  10  per  cent  of 
'•■'  cases.  Smaller  lirandics  may  li.>  occluded  without  any  evident 
'  I  iii!-'c  in  the  hcartlieat. 

■riicsc  results  iiidicale  that  the  capillary  areas  siijiplied  l>y  the  Itranchcs 
"'  '111'  coronary  artery  do  not  freely  anastomose  with  one  another.  They 
•I"   Mioic  or  less  terminal  arteries:  that  is,  each  liranch  supplies  a  distinct 

■■  -loll  of  the  cardiac  muscle.     If  oi f  the  smaller  liri lies  of  the  coro- 

■"  '   ''■'  •ludcd.  althoiijfh  there  is  no  immediate  s'oppa-fc  of  the  hearf- 

'  '  I,  .vet  after  some  time  the  area  supplied  l.y  that  l.raiicli  usually  iiiider- 
-    -  necrosis,  airain   indicating'   Ihat   collateral   circr.latioii  can   not   have 

'      "" ^'iihlishcd.     ft  is  int«'rcstiii«:,  however,  to  note  in  this  connection 

''  .iiialoiiiic  studies  have  shown  that  a  certain  aimuint  of  anastomosis 
'  •-  occur  hctwceii  capillaries  of  dilTcrent   hranches,  although  it   is  evi- 

'.  l!oni  the  aliove  oliscrvations.  that  no  adei|uat .Ilateral  circulation 

•iHcs  cstaliiished  thioii'.'li  this  anastomosis. 


I    i 


>   !| 


i 
t 


PERFUSION  OP  HEART  OUTSIDE  THE   BODY 

'   "I'ler   that    the    Mood    supply   throU!.di    the   coronarv   arteries    mav 
I'i'it-ly  maintain  the  normal  nutrition  of  the  cardiac  "muscle,  certain 


l(i-J 


•nil:  ciuci  i.ATKiN  OF  Mil:  r.i.ooi* 


f    ' 


.•..iiditioMs  must  !»•  ruHillcil.  Tlu-  ifi-ou'iiitioii  "I'  llifsc  cnn.litioiis  li;  • 
liccii  .•iiM-oiiiplisli.'il  h\  iilisci\;itiiiiis  on  llic  ("xciscd  lit'iirt,  I'm-  it  lias  Imi n 
I'oiiMil  tli.it  it'  tlii'.v  ii'-,  nillillol  llic  iiiMiiiiualinii  liciiit  can  lie  made  t(i  Ix  .  ' 
in  iiiil'i'ctlv  iiDiiiial  t'asliidii  for  several  Imiirs  after  its  reiimval  fi"' 
tlie  aiiiiiiai's  hiiily.  Indeed  certain  inaiiiiiialiaii  hearts,  siidi  as  tliat  nf  tl 
raliliit,  may  he  made  to  heat  for  several  days  outside  tiie  hody.  We  iii:i  - 
consider  the  essential  conditions  of  the  hlood  su|)|)ly  under  four  hi-adinu-. 
(  li   the  teiniieratnre:  i  •_' i    ihe  oxyv'cii  supply:  CI'   the  pressure:  and   <  I 

the  elieiiiie,il  composition.     Si cssfiil  i)erfilsioii  iimy  he  performed  wit', 

arti(iei;ii  saline  solutions  i  e.  >s..  Loeke'sV  hut  it  is  simiilest  in  invcstiirati'ij; 
the  relative  iminntanee  of  the  ahove  conditions  to  start   the  heart   p'< 
fusion   with   delihriiiated   hlood. 

After  hi lini:  an  anesthcti/ed  animal,  such  ;is  a  ilou'  or  a  cat.  iiniM 

no  more  hlood  c,in  he  i-eiiioved,  the  hlood  is  <lelihriMated  and  (ilteii  I 
throuuh  yauze  to  remove  tlw  liloin.  The  thorax  of  the  ilead  animal  l^ 
then  ipiiekly  opened,  lijjratiires  pliiced  around  the  main  arteries  spriii'^'iii- 
from  the  arch  <d'  the  aorta,  a  cannula  with  its  end  pointinu'  toward  tl. 
hcirt  iiiseited  into  the  descending'  thoiaci-  aorta,  and  the  latter  'n! 
across  helow  the  point  of  insertion  of  the  cannula.  The  heart  is  i!m  n 
ipii.'Uly  removed  from  the  thorax  and  an  artilicial  saline  solu  m 
(l.ocl-e'si   allowi'd  to  run   into  the  aortic  cannula   throiiirh  a  side  tnl.". 

until  all  thi'  hi I  has  heeii  washed  out   from  the  coronary  vessels.     Dur 

inir  this  operation  the  iieart  may  di'vclo])  a  few  heats  even  t'loiiyh  tic 
solution  is  (piite  cool.     The  aoitic  cannula  is  now  coiineeled  witli  a  hot'  " 

ciiilainiin.'    the    delihrinated    Idood    ililnted    with    I Ue's    solulicni    ;ii '1 

hroii'.'lil  to  hody  temperature  hy  imir.ersicoi  in  a  water-h;ith.  I'.y  nie;;M^ 
of  a  siiitahly  rcirnlatcd  air  presMiie  exerted  on  the  surface  of  the  dilii'  '1 
lilood  in  the  hottle.  this  is  forced  throunh  an  outlet  at  the  foot  of  i  !■■ 
hottle  into  tuhinii  whiidi  nins  to  the  .-Kntic  .'amiiila.  The  llnid  thus  li:  U 
its  way  into  Ih.'  coronary  vessels:  for  in  passinu'  toward  the  heart  in  "" 
aorta  it  will  <dose  the  semilunar  valves  iind  force  its  way  under  pres~^  ■<• 
into  the  coronary  vessels,  suhseipiently  escapin^r  hy  the  coronary  sinus  r  " 
the  riuht  ;iuri<'li'.  Very  soon  after  the  perfusion  is  started  the  li.  i 
lieiiins  to  heat  vigorously  and  reiriilarly.  thus  olTerino;  a  suitahle  prep-  ) 
tion  upon  •vhieh  to  test  the  first  three  mentioned  comliticnis  neees-  -.^ 
f.i,-  the  nutrition  of  the  cardiac  musculature  (  i-'i-r,  :!!•). 

If  the  h  ,iii>i  nihil)  of  the  solution  is  allowed  to  fall  considerahly,  'n' 
hciit  liecomes  milch  slower,  and  if  the  eoolini:  is  proceeded  with,  the  I  it 
will  iit'tcr  a   while  cease  heatiny;  alto<_'elher.      If  the  iinssiin    is  h.u.       1. 

the  heat   will  not   neeessaiily  h ime  slower  hut  very  much    feehlcr,       il 

will  sodii  cease.  In  Lieiicral  it  may  he  said  that  the  ienipei  atlifc  t,;  ^' 
v,,,lution  alVects  the  rale  of  the  heat,  and  the  inessure  alTects  its  stre'      li 


»;:      i 


Tin;  \  I  TUITION  (IP  Tin:  iiivirr 


lf)3 


Funnel 


Stock  fiftlon 
(Dilutea  iloodi- 

a  salt  solution) 
Metal  pan 
Hot  water  bath 


i^'     ■''■""'     I'Mii    ,.1    .,||.ii.ilii~    iHi     r,.,,,.iiiii!    tr,iiii;i;~    h..iii    .,ii    ,  vu-r.l    li.ait    I  I..11H!,  11.I..1  iT 

''        ""     '."■'"    1-    l-'l''    "■i""    I'N     ■!    i--il'i    I'-illi    lli..nl    u.iiTuTi.    an.l    111.     |.,rlu-i"n    lliii.l    i, 

"■'"""'        ""■    'iiiwiii,.    |,i.^v,,i,     III    this    ;i|.i.;ii.iiu-    1-    ^;i|.|.|ii,l    h\     ur.uilv.      ll-r.iTii    J.i.kM.nl 


H 

^^^IV] 

' }  '^^R' 

\'      s 


ir>4 


Tin;  ciitcn.ATHiN  of   iiii;  ui.iiiin 


(' 


I  H 


!.        I 


It  is.  iK.wfv.T.  ol.vimis  lli,it  :ii  ixTtuscl  i)i('));iriiti<nis  cliaiidi's  in  prcs 
sine  air  iik.-iy  to  ciiusf  alliMatinns  in  vat."  as  well  as  in  foicc.  unless 
•.'irat  can'  is  tak.-n  to  i<«'ci)  tlic  lioait  itself  as  warm  as  the  perfusion 
llui<l. 

The  iiiiimi-tanee  of  an  a<le(|uate  ))ressuiT  in  the  eoronarv  vessels  ha- 
heen  eleailv  hnniirht  out  in  eertain  exiMM-inieiits  in  whieli  the  heat  has 
l)een  maintained  for  a  shmt  time  hy  estalilishinj.'  a  pressnre  in  the  eor 
onary  vessels  by  means  of  indifTerent  fluids  or  <iases.  Thus,  if  oxyjien 
■las  is  allowed  to  pass  tlirout:h  the  vessels  under  pressure,  the  heart  will 
heat  for  a  short  time,  and  the  same  result  has  heen  observed  even  when 
mineral  oil  w  mereury  has  heen  i)erfused  under  press\ire  (Sollmaini ). 

The  necessity  for  an   ntliqiinii    oriiijiii  siipi)!!/  is  very   rea<lily  demon 
strated.     If  the  darker  hi.x.d  e.jeeted   from  the   litrht   anriele  with   eaeh 
heartbeat   is  transferred   ininn'diately  to  the  i)erfnsion  bottle,  the  heart- 
beat   will   s I    beccnu"    feeble   ami    irr<>!.'ular.   to    be    readily    rest<ired    to 

normal   when    this  dark    bhxxl    is  shalien    n)>   with   air   or   oxygen. 

Hy  artilieial  |)eifnsion  in  the  manner  above  described,  the  automatism 
of  the  heart  may  be  restored  many  hours  after  death.     Partial   restora 
tion.  •■onfine.l   to  the  aurieles  nr  to  that   part  (.f  the  venlrieles  lyinu:  iiii 
iiK'uiately  a.l.jaeent   to  the  lar^'e  blood  vessels,  ean  also  be  aeeomi)lisliecl 
in   the  heart    of  man  several   days  after  deatli.   ))rovided   death  has  not 
been  caused  by  some  acute  toxic  infection  snch  as  diphtheria  or  septic, 
mia.     The  Russian  physiolo-iist  Kuliabko,  has  succeeded  in  restorinjr  f"' 
over  an   houi-  the  normal   beat   of  the  heart   of  a   three-nn.nths-old   lio.\ 
twenty  hours   after  death    from   double   pneininmia.   btit    here   a<rain   tin 
indsation  returns  .ndy  in  certain  parts  of  the  heart.     As  will  lie  pointe.l 
o\it.  the  remarkable   resistan.-e  of  the   heait    muscle  displayed    in   Ihes- 
experiments  has  been   taken   as  an    argument    in    favor  of  the    my<i«reiiir 
hypothesis   for  automatic   rhythmic   power  of  cardiac   muscle,  the  armi 
mcnt   bein<.'  that   nervous  strnctni'cs  could   not    live  so  long  a   tinn-  afl<  : 
death.     The   fallacies  in   this  aryunu'nt    are  diMMisscd  elsewhere. 


RESUSCITATION  OF  THE  HEART  IN  SITU 

A  suitable  intracoroiiary  prcssun'  is  a  sim  <iiiii  mni  for  the  maini' 
nance  of  the  heartbeat,  and  this  is  a  fact  of  ..'real  clinical  siftniiican.- 
for  it   inilicatcs  that  any  attem|>ts  t.i  resuscitate  a  dead  animal  are  c  ■ 

tain  of  failure  unless  the  th.i.l  is  swh  as  will  biinu;  a   nutrient   llni 

under  a  certain  ))ressure  to  bi'ar  .)n  the  coromiry  arteries.  Injection  ■ 
fluid,  even  of  delibriiiated  blooil.  into  a  n  in  will  olivionsly  fail  to  U. 
(ill  this  condition,  for  the  )i.Mfnsion  must  be  nnnlc  into  an  aiter\-  s.i  ih. 
the  fluid  is  i-arried  ilown  \\u-  aorta  ami  tlienc.'  into  the  coronary  art.'ri' 


^ 


■rui':  N I  TUITION  (IK  Tin:  iii:.\kt 


16.1 


Till'  iiracfical  (|iicsti()ii,  in  so  far  as  resuscitation  of  the  licaitheat  i 
inr  nil)   irr  i/il  llir   iircf smirn  fluid   utuhr  itro 


ticcnicd,  is  tlicri'ioi'c.  // 


siin  into  the  hi niinihiii  of  lln  iiorla.'  Kvcii  if  we  were  to  transfuse  tliiiil 
under  eonsideraltle  pressure  into  the  aorta  throuj.'h  the  earotid  artery, 
il  uduhl  mainly  follow  the  iartre  vessels  leadin;^  away  from  the  heart. 
Hilly  a  fraction  of  it  icaeliin-r  the  lie-rinnin!,'  of  the  aorta.  To  "omi)eI  the 
iliiid  to  pass  t<n\ards  tlie  heart  we  must  iiitrodiu'c  some  obstruction  to 
Its  ])assiifje  perii)herally.  This  can  he  done  liy  the  injection  of  a  consid- 
.■i:ililc  dose  of  epinephrine  ( adrenaline  i  in  normal  saline  solution  throufrh 
I  he  needle  of  a  hypodermic  syringe  iusei'ted  into  the  tuhius  Ieadin<r 
iioiii  the  burette  or  pressure  bottle  to  the  cannula  in  the  earotid  artery. 
.\s  the  i)erfusion  fluid  is  running  in,  the  epinephrine  injection  is  quickly 
iiiMcIc,  artificial  respiration  and  cardiac  massaiie  bein;;  meanwhile  prac- 
liceil.  In  the  majority  of  animals  it  will  be  found  that  complete  res- 
toration id"  the  normal  blood  pressure  can   be  efVected  by  this  nu'thod. 

bi'l 1  liy  performinjr  the   resuscitation   under  aseptic  conditions,  some 

.iiiiiiiais  may  be  permanently  ifsuscitated  so  far  as  the  circulation  is 
■iiiK'crncd,  althous;h  the  nervous  structures,  even  after  a  few  minutes 
"f  •'death."  never  reac(|uire  theii'  normal  condition. 

The  epinephrine  acts  mainly  by  const iietin<;  the  snndl  arterioles  and 
'iiiis  directiii>r  the  bloodllow  to\\iii<ls  the  heart,  but  i»artly  also  by  a  direct 

>iiMnilatiM<{  acti the  cardiac  muscle.      It   does   not,  however,  cou- 

•!;ict  the  coronary  vi-ssels:  on  the  contrary,  it  is  said  to  cause  these 
-iiL'hllv  to  dilate. 


THE  RELATIVE  IMPORTANCE  OP  THE  VARIOUS  CONSTITUENTS 
OF  THE  PERFUSION  FLUID 

Hi'  can  slnd\  the  cheniicai  conditions  necessary  for  ifsuscitatioii 
'  ilic  heartbeat  by  obser\  inn  lli<'  beat  of  an  artiticially  perfused  heart 
lull'  solutions  of  ditTerent  chcTnical  composition  are  beiiifr  perfus'd 
"loiiirli  the  coronary  ve.s.sels.  At  the  (uitset  we  are  impiessed  with  the 
!'t  that  for  successful  resuscitation  the  or^'anie  c(nistitiu>nts  of  the 
i'liciit  fluid  are  of  trivial  impcntance  comiiared  with  th-  iMor<;anic 
■iistilncnts.     With  a  solution  containin^r  the  proper  iniipoi-tion   of  in- 

■.;aiiic  salts,  and  id'  eiuirse  an   ade(iuate  supply   o\'  o\\jren,  tin'   heart 

■I  rabbit,  for  example,  nmy  be  made  to  continue  beafintf  for  scveial 

'.^•^      It   is  true  that  it   will  beat   lonjrer  if  some  of  the  orsranic  eon- 

"iciils  of  the  blood  i)lasma.  particularly  carlxdiydrate,  are  present. 
•'    "II    the    inoriraiiic    constituents    alone    its    ability    to    beat    is    truly 

-■likable. 


3j^ 


\i    J 


166 


I  i 


■f 


Tin:  ciiicri.ATioN  of  Tin;  hi.ih)!) 
Observations  on  Cold-Blooded  Heart 


Tlic  ciirlicf  fxiu'iiipiciits  t'tir  tlic  iiiv«'stii.'iiti(m  of  llic  clicnrH-nl  citiiili 
lions  iii'ccs^jiiy  I'tir  llic  in;iiii1ciiiiii(M'  of  llic  licjirtliciit  wi'i'f  pci'fKiinc.l 
(Ml  //"  li'di-l  of  till  friiii  III-  htrllt.  r>y  iM'rnisiiiu:  cillicr  of  lliosc  licaft> 
^^itll  pli.v  >ic>liiirii'  sixlitiiiii'liluriili'  soliitinii,  it  was  (ilisci'vi'tl  lliiit  llimijili 
the  licnt  tniu'lit  contiiim'  I'm-  smiu'  liini',  >  ct  it  <j;ra(liially  -frcw  fccbliT 
aii<l  iVcIiliT,  iiiilil  at  last  it  ceased  allDiretiier  with  tlie  lieart  iniisele 
ill  a  eiiiKJitidii  of  I'xtreiiie  tela\alii>ii  or  diastole.  II'  small  jn'oportioiis 
of  ))(plMssiiiiii  and  cahMiliii  salts  las  cliloridei   were  added  to  tlie  sodium 

(•lilori<le  soliili the  heat  was  iiiiieh  hetler  maintained.     Doctor  Sidnex 

jiinsici-  i(i-o\cd  that  the  optimum  eoncentration  to  i)i-odiice  eflicient  and 
ItroloMircd  contraction  for  the  heart  of  the  tVo;;  or  terrapin  is  as  follows: 
jmtassium  (diloriile.  (».(»:>  jier  cent;  calcium  chloride,  (t.O'Jf)  i)er  eeiit 
The  ell'ectiveness  of  the  solution  was  also  loiiiid  to  he  increased  hy  th> 
addition  ol'  O.du:;  per  cent  of  sodium  hicarhonate.  Tliis  acts  as  a  hiil' 
(i-v  suhstniice  ( paj^e  :!(>).  lioldiMjr  the  h.\  dro^'cn-ion  concentration  at  :i 
constant  level.  .More  recent  work  has  sIkiwii  that  the  hydro'_'eii-ion  cop 
centratioii  ot"  the  ix'ifusioii  solutions  is  of  coiisiderahle  importance  in 
deterniinin<:  the  cfliciency  of  t!ic  heal,  lint  tlie  optimum  is  not  tlie  same 
for  the  hearts  of  dilTerciit  kinds  of  animal,  and  indeed  it  may  differ 
for  dilTereiit   parts  of  the  same  heart. 

T'u'  (lucstion  naturally  arises  as  to  the  relative  importance  of  eacl. 
(d"  the  aliove  salts:  or  rather,  \\f  should  sa.v .  cations,  since  the  anion, 
chlorine,  is  the  s;ime  for  all  of  them.  The  function  of  the  kixUiiiii  clihi 
lide  in  the  solutions  is  twofold:  ili  to  endow  the  solution  with  tin 
proper  osiiiotic  [ircssure  I  see  pa^re  4);  and  (2)  to  perform  the  spcci.il 
role  of  the  sodiiiiii  ion  in  the  oriyinatioii  and  iiiaiiiteiiance  of  llie  aiitn 
iiiatic  heat.  The  hitter  function  of  Xa  can  lie  shown  hy  oliserviii'r  tlie  lieliav 
ior  of  strips  cut  out  from  the  vciilricle  of  the  turtle  heart  and  idacc! 
in  siilulions  of  correct  osmotic  iiressure  hut  coiitaininji  no  sodium  chh' 
ride  isotonic  soliilions  of  cane  suj;ar,  for  esample.  Tiiey  soon  cea^' 
to  heat,  hut   if  a  y  loiiiit   of  sodium  idiloride  is  adiletl  to  the  cam 

sinrar  solution,  rhyii..  ..■  ci-ntractioiis  return.  The  role  of  the  riilnin, 
inns  is  almost  eiilirel>  a  iiharmacoloyic  one.  If  a  strip  of  turtle  ven 
tricle  which  has  lieeii  made  to  cease  heatiiii,'  liy  immersion  in  isotoni' 
siijrar  solution  is  placed  in  a  weak  solution  of  calcium  chloride  hefor 
it  is  traiisferre<l  to  sodium  chloride  solution,  the  si»ontaiieous  contrar 
lions  will  return  earlier  and  continue  for  a  hmirer  time.  On  tlie  otln 
hand,  if  more  than  the  c<irrect  amount  of  calcium  salt  is  present  in  ll 

.solution,  the   heats  will   s i   lie    found   to   liecoiiu-  siualltM'  and  small' 

in  amplitude,  hecaiise  i-ela\ation  does  not  i^-ojierly  occur  hetween  then 
and  ultiinatelv  thev  will  cease  altotrelher  with  the  ventricle  in  a  coiiditi'  i. 


,\- 


TlIK    .MTKITION"   OK   Till;    III.AKT 


167 


I'xti'cuic  (•(iiiti-iiftioii,  ciilli'tl  ciilciuni  i'i«(ir.    The  inijxiftancp  of  rdlclum 
y  iilso  lie  slidwii  l>y  iitt(<iii|>tiii<;  to  pcit'iisc  a  lurtlo  licart   with  Mood 


t!i   from   wliicli   the  calcium  has  h 


removed  bv  the  aildiliou  of 


s..iliiiiii  oxalate  (which  precipitates  it  as  iiisoluhle  calcium  oxalate).  The 
liiMit  soon  ceases  to  l)eat,  liut  can  readily  he  math'  to  do  so  ajjain  by 
jiildiii!;  a  slifiht  excess  of  calcium  chloride. 

The  potdnsium  ions  <h)  not  ai)i)ear,  like  those  of  calcium  and  sodiuin,  to 
!"•  alisolutely  essential  for  the  iiiaintenaiice  of  the  liearthcat ;  at  least  the 
liiiirt  of  the  turtle  will  beat  for  a  loii'r  time  when  perfused  with  a  solu- 
linii  cnntaiiMu;.'  only  sodium  and  calcium  salts.  The  explanation  of  this 
ifMilt  need  not,  however,  necessarily  be  that  potassium  is  an  unessential 
rnii>tituent  of  the  perfusion  fluid,  for  it  may  well  (h-pend  on  tlie  fact  lliat 
ihiic  is  a  suflicient  store  of  ]iotassium  locherl  away  in  the  muscle  fiber 
111  siip|)ly  the  re(|uirenicnfs  of  the  heart  muscle  for  tiiis  ion  foi'  at  h'ast 
;is  loiu;  as  the  beat  would  continue  under  any  circumstances.  In  any 
r:is(',  we  know  that  potassium  has  a  i)rofound  inllueneo  on  the  lieart- 
li.iil,  for  when  the  jiroportion  of  it  in  the  pi'r*"usion  tluid  is  inci-eascd.  the 
In  .il  hecornes  very  slow  and  the  tone  of  the  heart  is  <rreatly  diminished — 
t!  ,;i  is,  if  becomes  extremely  relaxed  between  the  beats;  and  if  the 
;iiii'iiiiit   is  further  increased,  will   very  soon  come  to  a  standstill   in  a 

•M.itly  <1ilated  or  diastolic  i)osition. 

The  '<trikin<;  antat;onism  displaye(l  liy  these  inorfianic  cations  upon 
i' r  lieaitbeat  has  led  some  iiivestij,'ators  to  su<;j,'est  that  the  stimulus  re- 
S'i'iisible  for  the  rhythmic  activity  of  the  heart  depends  on  some  sort 

■!  I'liiiiiical  union  oceurriny;  between  tlu'  inori;anic  cations  and  the  con- 
!  ■■liile  sidisfance  of  the  heart.  I'liion  of  calcium  witli  the  contractile 
^  stance  will  lead  to  systole  or  contraclioii.  w  heicas  union  of  sodium 
I  ■   |iiitassium  will  lejid  to  relaxation  or  diastole. 


■  ■■! 


Observations  on  Mammalian  Heart 

liivesti!;ation  of  the  efticiency  of  various  saline  solutions  on  the  iso- 
■i!  nKiiiniKilidii  Ik  art  has  shown  that  the  iu'o))ortion  of  the  ■diove  s.'lts 
-t  lie  somewhat  dilTcrent   from  that  used  for  the  cold-blooded  lieai'f. 

iiiiuht  be  <'\peeted.  the  most  etlicient  projiortions  are  those  ])rcsent 
'\\v  blood  serum  of  the  i»articulai-  animal  whose  heart  is  bein^  jjcr- 
'  1.     Hasin^  his  |)ro|)ortions  upon  the  results  of  analxscs  of  the  iiior- 

!!•  ciiiistituents  of  mammalian  blood  serum.  liOeke  found  that  an 
".iiuiic  solution  of  the  following;  comi)ositioii  is  most  etlicient:  so- 
:iii  chloride.  (),!)  per  cent;  cisicium  chloride,  0.024  ]i(>r  cent;  potassiuiTi 
'  'lide.  0.042  per  cent:  and  sodium  bicarbonate,  O.Ol  to  0.0:>  per  cent. 
'  !  II  "liocke's  solufi<»n."  as  it  is  calleil.  is  jierfused.  with  oxy>;en  in  it, 

111-  pressure  through  the  isolated  mammalian  heart  at  body  tempera- 


1G8 


Tin;  ciui  I  i.ATKiN  (IF  Till;  iii.oon 


r  I 


till'!',  clllicicnt  ln'iitin^r  fiin  lie  iiiaintaiiicil  i'lpr  iiiiiiiv  luiiirs.  Jloi'f  ri'ct'iitl 
a  soliitiiiri  known  <is  Tvi'dtlc's  Ik  (■iiiiinionlv  used.  It  cinitiiiiis  a  small  aiiiiiiiii 
(if  iiiaL.Mi('Niiiiii  and  of  pli<is|iliat*'s.  Altliiiu<:li  iindiiiililcdlv  supciiiir  I'" 
sdiiic  jici't'iisi'd  |irc|iaiaticins,  siii-Ii  as  liic  intestine,  it  dues  imt  seem  tn  I- 
ill  any  wav  siiperidi'  tii  l,iieke"s  t'cn-  tlie  pei-fiisi<iii  (if  the  lieart.  'I'lie  liiiMi 
Inmates  and  pliospjiates  in  tliesi'  siiliitinns  endow  them  with  a  hvdro^en-ii';! 
eoiieentration  near  tliat  of  the  hlood  (slisrlitly  on  the  alkaline  side  ni 
neutrality',  and  at   the  same  tMiie  they  act   as  liiilfef  snlistaiices. 

As  ;dii'ad\  pointed  out.  the  oriranie  constituents  of  such  iierfiisinri 
lliiids  i|o  not  appear  to  he  relatively  of  nearlv  so  nnndi  im|)ortaiiee  .>- 
tlie  inoifranic.  Nevertheless  it  appears  thjit  a  small  pereentafre  nid! 
lier  cent)  of  Kliicose  does  materially  iinpro\e  the  nutritive  (pialities  o: 
the  solution,  and  it  has  moreover  lieeu  shown  tli:it  after  a  while  the  eipn 
eentratioit  of  >;lneose  in  the  perfusion  llni<l  distinctly  decreases.  Tlr> 
does  not  of  itself  necessarily  mean  that  the  (ilncose  is  actually  iitili/r.| 
liy  the  heart   muscle:   it   mijiht    he  stored  away   in   it   as   irlyeojien.     Tli.-  ; 

some  consumption  of  carhohydrate  does  however  •nr  in  tlio  lieart  h:  - 

lieeli  demonstrated  liy  measuriii;:  the  intake  of  oxv  treii  and  tlie  oiitpiii 
of  cailion  (lioxidc  fliroilyh  the  luntrs  of  an  isolated  heart-lunjr  prepai; 
tioii  |)crfnscd  outside  the  hody  with  defilirinatcd  Mood.  P.y  experiiueiits  ..l 
this  type  the  attemjit  has  heen  made  to  show  that  the  heart  of  dialiel 
animals  loses  the  power  of  linnuiiur  <;lncose  ;is  compared  with  the  liearN 
of  normal  animals.  While  the  cxperiinents  arc  \eiy  sii};^estive,  tin 
results  do  not  as  yet  justify  lis  in  claiiuiiiir  that  in  the  latter  disease  t!  • 
power  of  liiirninu:  glucose  in   the  tissues  has  liceii   materially  depressil 

Tlir  ioiicfiitriilii)ii   of  hiiilnifit  II   iins   In   llu    iiirfiision    fluid  has  an    iii. 

portant    influence   on   cardi; Ilicicncy.      We  also  know   that    the  iim    i 

coiivciiieiit   iiiethod   fir  chaiiiiiiii;  the  hydroiren-ion  eoiieentration  of  sii.   - 
llnids  is   liy   ,dterin<r  their  tension   of  eailion   ilioxidc    (s<'e   pafre  ;!.")4l.      !'i 
a  heart-liiii!.  pi-'paration,*  such  alteration  in  carlion-dioxide  tension  r.   , 
very  readily  lie  liroiiclit  aliont  liy  alterinir  the  pcTceiilaj;o  of  this  ;ras     < 
the  air  with  which  the  luntrs  are  ventilated.     To  nieasiiro  the  efficici 
of  the  heartlieat   in  such   an  experiment,   it   is  convenient   to  enclose  ' 
oi<iaii   in  a  cardioplethysmo<rraph.  the  tracing  of  wliicli   will   tell   us  ' 
dcfrrce  to  which  the  heart  is  eonliacted  or  relaxed,  as  well  as  the  outi    < 
of  hlood    per  iiiinntc.      My    inereasinj;   the   tension    of  carlion   dioxide,     i 
has  heen  found  in  such  experiments  that  the  dilatation  of  the  veiitr:    ■ 
is  oneoura'^eil,  so  that  the  heart  with  each  heat  diseliarsres  a  liiryrer  (pi     • 
tity  of  lilood  (I-'i<j.  4Hi.     When  detilirinateij  Mood  is  used  the  optiin    ■' 


•.■\   lir.irt  IiiiiR    itcpiu.tii" 
thr    vii-^rN   hciiiy    --'Mtjltly    i< 


in    \\lii.  1i    I.. .ill    )i(.(it    an. I    liinu-    .in     t.tiln-t.l 

'U'l  to  inaiiMuii!  .1  I  Mtitiminti^  « in  iil;tiiiiii 


illM.I.    111. 


■illi;  M  TUITION  OK  Tin;  iii;aiii 


1G!» 


|.n 


sMirc  III'  Ifiisioii  ol"  cnilnni  dioxide  liiis  liccii    fomid  ti 


1   lie  lii'twccn  ■! 


(I  to  ])( 


r  iM-nf  of  tin  iitiiiosplicrc. 


Tllilt  tlic  ctTt'ct  of  cjirlioii  dioxidr  in  en 


<-our'ii<;in<;  tlic  rchistitioii  o 


f  111 


•ait  Ixtwccii  liciits  is  ilcpciiilciit  upon  tin'  cliaiiy;!'  in  liydro<^on-ii)U  con- 
iitiatioii  of  the  p.-rfiisioii  lliiid  lias  Imh-ii  sliowii  bv 


ivsiilts  ill  cxpcrinu'iits  with  i)crlusion  lliiids  tc  wliidi  iJilTcirnt  ipiaiit 
lilies  of  weak  lioiivolatile  acids  lia\ 


seeiiriiij;  tlic  same 


e  iteeii  aildec 


Idi 


TI 


lese  ol)sei'\atioiis  are 


'■"■'v;"''".^"'""'f   ''"'"'■   "f   v.-i.tnil.s   „f  ,at    (l.,w<r   .,im.)    i„    a    li,  arlliniit   perfusion    prepara- 
";       llK-air    llsnl    til   viMtilalc    the    liinns    was    riplaiiil    iMlwi.n    the    arrows    l)V    a    mixture    con- 

i.-tmg  .11  ,    to,  ail. 1   :y',    (),.     Thu  lau^.-.l   ililaiali jf   ih,    v.iilricl.s  aloiig   with   fcelilcr   beats 

'.1    a    teii.lcmy    for    the    arterial    pressure    to    fall     (upper    eiir-.el.       The    after    effect    was    an    ini- 
•iveiiient    of    the    heat.      (I'rom    SlarlinK.) 


'I   praetieal   iiii|ioitaiiee  lieeailse   of  the   liy-lit    wliieli   lliey   tliniw   on   tlie 
Miise  of  cardiac   failure   follow  iiijr  upon   conditions   in   wliieli   there  lias 

' II  excessive   removal   of  cailion  dioxide   from   the  Mood,   as  in   forced 

•'iitiiation   of  the   luiif;s.      Vaiidcll    Henderson   has   su};<;este<l   that    siir- 

L'';il  shock  may  lie,  partly  at  least,  due  t<.  cardiac  failure  followiiij;  the 

wishiiii:  out"  of  earlion    dioxide    fiom    the   hlood    hy   the   dysiiiiea   so 

ten  incident  to  the  adniinistiati ,f  anesthetics  in  sul•^'ical  ojieratioiis. 


jl^' 


ifl 


1.1 


CIIAl'TKU  X.\ 

Till-:  riiYsi(iL(MiY  <)i'  Till-:  iiKArrrMKAT 

THE  ORIGIN  AND  PROPAGATION  OF  THE  BEAT— THE  PHYSIO 
LOGIC  CHARACTERISTICS  OF  CARDIAC  MUSCLE 

The  ima'iu  and  |)iu|)iit:atiim  oftlif  lit-jittlx'af  air  studied  mi  flic  i-xciMil 
licait  of  a  t'riii:  nr  tiiitlc.  or  on  tlic  inaniinalian  lit-art  by  j>i'it'usin}{  it 
undiT  snitalilc  conditions,  wliicli  have  alifady  lii'cn  dt'scriln-d.  The  result^ 
olitainci!  on  tlic  cold-liloodcd  Iicart  apply  H'oic  or  less  diicctly  to  tln' 
^val■nl-lllood(•d.  In  tlic  first  i)la('c  it  is  i-lcar  that  tlic  rliytliniic  contrac 
tility  of  the  licait  is  not  at  all  dc|icndciit  upon  tlic  central  nervous  svs 
teiii,  for  il'  it  were  so,  the  excised  heart  could  not  contiiiiie  licatinjr.  Tlii- 
fact  docs  not,  however,  necessarily  imply  that  the  licatini,'  power  is  in 
dcjieiideiit  of  in'ivoiis  st rilctlires,  for  in  the  heart  itself  an  extended  net 
work  of  nerve  cells  and  coniicctiii";  nerve  fillers  can  readily  he  denion 
strafed.  It  iiiiylit  (piife  well  he  the  case  that  the  rhythmic  beat  is  dc 
pendent  upon  the  transiiiission  to  the  muscle  fibers  of  the  heart  oi 
iiiil>lllscs  ircncratcd  in  the  ner\c  cells  and  transmitted  aloii};  the  nei\.' 
fibers  of  this  local  nervous  system.  Siicli  is  the  ii<  Kni'/i  nlf  liiifxiihi xis  i,! 
till    hcnrthtiil. 

On  the  other  hand,  it  may  be  that  these  nervous  structures  are  not  nl 
all  res|ioiisible  for  the  orii.'ination  of  the  beat,  but  serve  merely  as  st;i 
tioiis  on  the  piitli\\a\-  of  the  ncr\e  impulses,  transmitted  to  the  licai' 
from  the  central  nervous  system  aloni;  the  vai;us  and  sympathetic  iiervi  -, 
for  the  imrpose  of  alleriii<;  the  rate  of  the  heartbeat  so  as  to  ad.just  it 
to  liie  re<iuiremcnts  of  blood  supply  in  the  various  parts  of  the  body.  I  i 
siiidi  a  i-ase  the  rhylhniic  ])o\\cr  would  reside  in  the  iiiuseidar  tissues  '  i 
the  heart  that  is,  eaidi  cardiac  muscular  cell  would  have  the  jiowi 
not  merely  like  skeletal  muscle  of  coiit raetinj,'  ill  resiionse  to  a  stilllu^i^ 
traiisiiiilled  to  it.  but  also  of  (oitrinat injr  that  stimulus  within  itsc  ;. 
This  is  the  iiifiiiiii  nil-  Iniiiolhatis.  .Much  controversy  lias  rafied  arou'  1 
these  two  hypotheses  and  altlioii^rli  space  will  not  iiermit  a  detailed  stii  . 
of  the  ipiestion.  it  will  be  ni'ccssary.  on  account  of  the  jrreat  iiiiporfaiiiT  ■! 
the  siib.ji'ct  from  the  physiolojric  stand|)oint,  briefly  to  review  the  in.  " 
aiirniiicnls  of  eacii  scliool  of  thouu'ht. 

There  is  no  piece  of  evidence  ofTcred  by  the  advocates  of  either  '    ■ 
iieiiroi:enic  or  the  niyoy;i'nic  hyiiothesis  that   can.  taken  singly,  be  <•<   :■ 

170 


I 


TIIK    PlIYSIOI.CMiY    (»F   Tin:    IIKAKTIIKAT 


171 


-i.liit'd  as  iil(s(ilnl<'ly  fimclusivf.  Altliouirli  sonu"  of  "tlic  jx'nnfs"  iiiiiy 
;it  lirst  Niy;lit  iipix'iir  to  !)(•  (••iiiclusivf,  jft  cacli  of  tlicin  Incnks  down  when 
>u|pjc('t(M|  to  II  closer  siM'Utiiiy.  It  is  only  iil'tcr  we  liavf  ■.•ollt'ctcd  nil  the 
(Uili'iicc  for  jind  iiirainst  cacli  view  that  \vc  shall  Itc  in  a  position  to  conio 
III  any  coiK-lusion,  and  even  tiicn  it  will  he  i>lain  that  out'  coni-liision  can 
lir  iiiily  tcntativ*'. 

Myogenic  Hjrpothesis 

Takinir  first  of  all  the  evidence  in  support  of  the  niyo)j;i'iiic  liypotliesis, 
till'  follouiii};  stands  out  most  ]>roniinently: 

I.  The  heart  heats  in  the  end)ryo  eliiek  Itefore  any  nerve  cells  have 
L'liiun  into  it,  and  not  only  tliis,  liut  if  jxirtions  of  heart  muscle  are  re- 
imivrd  from  the  endiryo  and  i)laced  in  blood  plasma,  they  will  continue 
I'l'Mlirn.'  fill'  niaii.N'  days  It  lias  also  Iieen  oliserved  that  cells  may  wander 
I'lT  from  this  mass  of  car-diac  muscle  and  undergo  multiplication  nnd 
(lilTiTciitiation,  so  as  to  produce  isolated  muscle  cells  whicdi  exhihit 
tii\lliniic  contractility.  Tiie  reltuttal  on  the  part  of  the  neurosrenists  of 
ilii>  apparently  iiiuissailalile  evidence  is  to  the  cITict  that,  althoU!.'h  em- 
lii\<iiiic  muscle  cells  may  exliiliit  the  power  of  rliythmii'  contraction,  this 
iliiis  not  mean  that  the  fully  dcveloju'd  muscle  cells  will  necessarily  luivo 
•^Ml•ll  |>o\\('r.  In  the  eary  stajres  of  emltryonic  devcloi)ment,  it  is  of  course 
(vi'leiit  tluit  tlie  functions  which  in  tlie  fully  (h'veloj)ed  animal  are  del- 
1  Liatcd  to  various  special  o'rtrans  and  tissues  should  he  ))erforMied  I)y  cells 
iiiniiiy:  several  sucli  functions  in  common.  The  muscle  cells  of  the  heart, 
<<iv  cxamiile,  nuiy  to  start  with  he  jiossesscd  of  a  power  rmt  oidy  of  con- 
I  itiuf;  hut  also  t>\'  initiatiu'.;  thi'  cont I'action.  It  may  he  that  tliey  are 
|i.!'!!y  nervous  in  character  and  that  only  later,  when  the  differentiation 
!■<  idiisudMuated,  does  the  i»ower  of  i-h.\thmic  contraction  become  (h'le- 
-■  it'll  to  the  nervous  elenu'nt  and  that  of  contraction  retained  by  tlie 
''I'l-i'lc  itself. 

-.  The  nervous  sti'ucture  in  the  heart  may  be  damaired  eithei'  by  me- 
(•'  nical  ineaiis  or  by  drujjs  without  apitarently  interferiii(i  with  the 
""  f  of  rhythmic  contraction;  for  example,  in  the  heart  of  lar«e  tur- 
•  '■N  it  is  possible  to  dissect  out  a  considerable  amount  of  nervous  tissue 
"  itlidut  any  disturbance  of  the  beat,  iiiid  in  all  animals  the  administration 
1  ^itropiiie,  wliich  ])araly/es  the  posti;an>;lionic  fibers  of  the  autonomic 
!      Mills  system  (sec  pajre  'J2*i)   found  in  the  lieart,  iloes  not  an'ect  it. 

The  ai)ex  of  the  ventrielp  in  such  hearts  as  that   of  the  turtle  can 

■!  invii.  by  careful  histologic  examination,  to  contain  no  nerve  cells, 

;  ■liiiioiij'h  a  few  nerve  tibors  may  be  foiuid,  these  are  functionless 

'lit  nerve  cells.     This  virtually  nerveless  ])iece  of  heart  mii.sele  can 

!ii.i<ic  to  contract  rhythniioally  by  perfusing  it  with  suitable  saline 


<,   ,- 


I7;i 


'I'lIK   I  IHcl  I.ATKIN    (IK    Till.    Itl.lH.H 


MtliiliiiM  iiiiiliM'  pi'i'sMii'i'  mill  >tiirliiitr  till'  licatinp;  lix   iipiiliciitinii  iiT  t'l 
ti'ii-.'il   >tiinuli.      iMiliitril   stiips  til'  vciitriiMiliir  iiiiisi-li',   in    \\lii<-li   iilso 


i  ^   I 


lU'lVolls   I'lfllli'llt    r.lM 


<l)'iiiiiM>tratfil.  iiiiiN    iiiiilrr  ravnnililc  cniKliti' 


Ik'  ctiiiscil  fii  lit'at  (|uili'  ri'iiiilarly  it'  sii|ipliiM|  with  pni|ici'  imti'iciil  llin 
Tlic  rrliiittal  of  this  cv  idfiu'c  is  twnt'iiM:  In  tlu'  liist  plarc,  skeletal  inn 
ele  itself  iiiider  eeftaiii  eoliilitiiilis.  such  as  exposure  to  solutions  en 
taiiiinu:  an  eseess  of  phosphate  i  Mii'ilennaiiii 'si,  may  exhihit  rhvtliin 
eontrartility,  esperially  on  eoulin^'.  wliieh  inilieales  that  exhiiiition  of  rli.M 
iiiie  power  in  isolateil  portions  of  eanliae  niiisele  iieetl  not  mean  that  iiml' 
oiiliiiai-.N  eonditioiis  sueli  power  is  responsilile  for  the  normal  heartltr;! 
Ill  the  se<  oiul  plai'e,  it  is  pointed  out  that  although  we  eaii  not  rev< 
their  presenee  liy  present-day  histolouie  methods,  this  is  not  eoiielusi 
e\  ideiiee  that  tlu'  heart-musele  tilier  may  not  possess  sonio  nervous  slih 
tiires  eapalile  of  fiinelioniliir  as  nerve  cells. 

The  heart  even  of  mamnials  can  lie  made  to  coiiliuiie  liealiiii;  for  ^. 
eral  <lays  after  excision  from  the  Imdy.     The  ner\e  cells,  as  we  know  tin  ■ 
in   (III'  ci'iitral   nei'xous  system  at   least,  can   not,  on   the  otlier  hand,   ' 
made  to  fiinctioiiale  for  more  than  a  few    hours  after  death.     Therefon 
it   is  ai'LMicd,  the  liearllieat   in  siirviviiiir  mammalian   hearts  can   not   ''■ 
peiid  on  the  IP  r\  oils  structures.     The  ar<;iiiiient  is  however  easily  refiiti 
on    the  one   hand,   we   do   not    know    that    the   nerve   structures   sitini!' 
peripheially  in  the  heart   muscles  are  of  the  same  viahle  nature  as  lli. - 
composing  the  ceiitrat   neiMius  system;  and,  on   the  other,  the  siirvi- 
of  the  heart  may  in  itself  lie  siitllcient  to  maintain  arinind  tlie  nerve  n 
emlirddcd  in  it  a  nutrient  cnxironment  ^\hieh  is  mneh  more  ))hysio|ci'  i 
than    that    which    we    can    snppl.v    in    artilicial    perfusions    of   siirviv  r 
nervous  tissues. 

4.   ( 'ireunistantial    hut    nevertheless   stroni;    evidence    is    furnished 
the   fact    that    many  other  \arieties  of  involuntary   mu.side  are   eiidm 
with   rhythniic  contractility;   thus,   the   muscle   of  the   intestines,   of 
ureters,   of  the   hladder.   of   the    uterus,   of   the   lilood    vessels   of   eei  t 
animals,  and  of  the  Ixiiiph   vessels  in  the  so-called  lymph  hearts,   m;   ' 
lain   rhythmic  <-outractility  .-iftcr  isolation   from   the  animal   hody.     'H 
rhythmic    ])oW('r  seems   in    certain    of  these   cases   to   he    independeiii 
nervous  control. 

Neurogenic  Hypothesis 

III   t'avor  of  this  hypothesis  the   I'ollowiliH:  evidence  is  offi'i'ed: 
1.  The  heart  of  certain  animals     id'  Liiiiiiliis,  the  kinn-erah,  for  ex    n 
pie,  is  dofiiiiiei.\    dcpeiideiit    i"!!!'  its  iliUlimic  contractility  uj)oii   n.     ' 
horiiifr  nervous  structures.     The  heart   of  this  animal   is  a  tiihiilar 
ciliated  oi'.Mii,  and  aloiiir  its  dorsal  surface  there  runs  loii<i;itiidiiia! 


Illi:    l'IIV>liil.iHa'    UK    Till     III   MITIIIXT 


171 


■J 

I 

1 


IP  i\i'  riii'il  iMiiihiiiiiiiu:  uiiiiuliiiii  i-i'lls  mill  iri\iiii;  nlT  IIIxtn  wliit-h  \u- ><(! 

ih  jiiii't  ilircctiv  til  till'  lii'tii't  mill  in  piii't  to  liitt'i'iil  i-i>i'i|s  i  Tiv.  41  >.  \{v- 
ihuval  III'  tliJM  iiKMliiin   iii'i'M'  iMinl   i^   t'liljuwctl    liv   tnttil   alinlitliiii   nf  tin' 

ii' altlirat  :   the   liriirt    I (Hiii's   |ii'ilVrtly   ijiiii'si-i'iit    liki'  iiii    iiiisliiiiiiliiti'il 

>Lrlftal  liiuscli'.  Ill  a|>|>l'ai^illl;  llif  i'\  iilriirr  at  its  tiiii-  xahic,  it  liillst  Ix' 
iimIiiI  that  althiiiii;li  liy  stiiniilatinii  ut'  the  iirr\<'  lllii'i>  i-iiiitrartimi  nf  tin' 
li.ait   ran    In-    prmlurril.    Ilir   rmit  I  artimi    i*    like    that    nf  a    sUflctal    iiiiin- 

il  tliiv  is  iiiiiNt   iinpiii'taiit     if 


it  is  lint   rlivtliiiiir :  ami  iimri'iiv  cr 
xai'iiills  plivsioliisrif  prnpci'tii's  nl'  iiiiisclf  as  <li'.si-rilii>il  lu'lnw    lie  stini 


I 


I  I  lap'    ITtil.  it    will   III'  t'liiliiil   that    in  all  nt'  thi'iii   thr  <|llii'H<'i-iit    lii-ait 
i,ii;m'Ii'  IicIiiivcs,  iiiif  like  the  lii'jiit  imisi'li'  nf  iitln'f  aiiiiiials.  Imt  like  that 
'  r  sUflrtal  iiiiisclc.     This  i'\  iilciiri'.  tlit'rcfnir,  whili'  iiiilispMtalply  shnwiiiL' 
iIimI  tile  heart  (if  Liiniiliis  ilcpi'inls   I'nr  it,  rh\tliiiiir  pnwn-  lipmi   ni'iu'h 
'.iiiii!;  iii'ixc  stnn-tiircH.  ilncs  imt  justify  tlu'  iissiiiiiptinii   that    this  \\ill 

I"   ll asc  ill  the  heart  nf  aiiiiiials  lia\  iiiu:  ililTerelit  pliysiiih>u'ii'  prnperlies. 

■J    The  ilispnsitinn  nf  the  liervniis  striii-tlires  in  the  heart,  espeeially  nf 
Mm-  iViii;  anil  turtle,  exaetlv    enrrespnliils  In  tiie  ileu'lee  nf  ilex  elnpiiiellt   nf 


-il       lit. tit    aiitl    t.ttili.tt'    lit  I  w,    t>l 
ill.    lutfiut    .-irtt-rk-K;    ht.    lulcral    iutvih 
.1.    riifh    |iatr    nf    whiili    itiirt'spttiiti^    Iti 
ittril. 


i.,niii 

n^      l-ni^liirtiiii^. 

(( 

;iiI-.Mi.  ) 

(III.     aiiti-rinr    ar 

ttnic, 

mcilian    K'lnx'i' 

nif 

Iiaiti;    Of. 

u'»lii    or   atTi-renl 

4)|K'      < 

t    the    si'^nit-tils 

iii1< 

uliiih    1 

If    l.imnlns    hi-art 

i'   itivllmiie  jinwer  nf  the  ililTereiit  parts  nf  the  heart:  tlius.  the  •rreatest 

'liinii'  pinver  is  iiiaiiifi'sted  In   the  sinus  ami  the  least  liy  the  tip  nf  the 

iiitiilc   at    the   liullius   arterinsus.      In    the    fnriiier   pnsitinn    the   nerve 

iHlilies  are  xery  ])riiininent  :  in  the  hitler,  iin  iierxe  ei'lls  ami  hut   few 

■  M-   lilicrs  eaii    lie   lieteeteil.      This    prnnf   is,    Innvever,    easily    assaileil. 

ll'  first   plaee.  it  may  merely  lie  a  eniiieiileiiee  that  the  ilispnsitinn  nf 

miM'  struetures  ami  the  (le\  eln])iiii'iit  nf  rhythiiiie  piiwer  i'nrres])(inil. 

'■   iiiii'(|ual    rhytlmiie    pnwers   may   ilepeml    primarily    nn    a    ililTerenee 

^    ^'mi-tiire   nf  the    iiuisele    filters   themselves,   siieli    ililTereiices    haviim: 

':■   slinwii    tn  exist    lietvvceii    the    nillsele   (•ells    nf   the    sinus   and    thnse 

-■!>,  the   xentriele.     The   fni'iner  cells,    fur  example,   have   much   less 

'  I'lpi'd   crnssed   striatinii   and    their   prntnpl.-ism    is   much    iiinre    u:iaii- 

iii  shnrt.  they  are  much  mnre  emli'yniiic  in  type  than  the  cells  frnm 

ip  nf  the  ventricle. 

'    .1  jury  had  tn  return  a  vei'dict  frnm  evidence  nf  sn  c(in(tictiinf  a  cliar- 
it  wiiiild  nil  dniilit  lie  eipiivaleut  tn  that  nf  the  Scnttish  cniiit    -■'tint 


;  I 


i! 


174 


Tin;  (11(11  i.ATfoN  ity  riii.  iiiJMir> 


Itfovfii,"  Hut  it  in  likflv  llmt  tin-  ii.ii.ji.iity  of  tlic  jmv  uoiiltl  \..i. 
in  fiiNor  of  llic  iiiyinr.iiii-  li.\  imlliois  rtnlnilily  tlif  safi-st  vicwpoini  t.. 
tiikc  lit  till-  iitcviil  tiini'  is  tliiit  tin'  |Ht\vcr  id"  ilivtliinic  I'lmf nicf inn  u 
inlHTi'iit   ill  tl iiiiliiii-  iiiiiM'Ic   (ilx'iH,  iM'iiiif  ni"st   liialilv    (IfM-lnpril   .m 

tlinsr  (if  till-  N.MI.MIS  Cllil  III'  llir  In-lit,  tilul  Icllsl  <1i'VI-|o1»1mI  ill  tllllSC  .  t' 
flic  lirti'lill!   rinl.      Sllrll  ll   r(>lli'lll>iii|l  iIiii'H   not    lirllX    to  tllc   IHTVons  Htiu 

Inns  of  llir  lii'iiit    ilic  powir  iiiulrr  i-crtiiiii  coiHlilioiis  of  iilso  nssmnii   ' 

rhvfliiiiii'  iirlivity.     In  •use  iit  li'ii^t      iiiiiiii'ly,  the  lirait  of  Liiiiulii^ 

«!•  know  tliiit  this  is  so.  l''of  some  rciison  in  this  aniniiil  flic  riinli^i  • 
imisc  Ir  liiirr  liiis  lost  its  iiiliiTi  III  ihyllmiii'  iiowrr.  ainl  is  now  ilfiinnl.  ni 
for  its  afti\ilifs  iijioii  iliylhiiiic  fivoiis  (lischaiiri's  f raiisiiiiftfil  to  il 
fioiii  the  nciu'lilioriii!.'  iiriM-  conls,  a  coiKlifion  which  is  jiaralldccl  m 
flic  hiu'hci-  animals  in  the  inncix  ation  of  the  rcspiiatofy  iiiiiscics.  TIm 
res  I  lira  ton  center  ili.v  Ihiiiiea  ;  discharu'cs  iiiipii'ses  lo  flic  iiinsi  Ics,  whi^  h 
are  (|iiiescent   in  llie  alisei f  these  ini)nilses. 

The  Pacemaker  of  the  Heart  and  Heart-block 

In  a   voliiiiie  of  this  nature,  devoted   primarily  to  the  practical   api'li 

cati f  jihysioloiry.  the  discussion  of  these  prolilciiis  may  seem  a  liltl'' 

old  of  placi'.  Iiiif  that  this  is  not  the  case  is  seen  when  we  consider  tliit 
the  experinieiits  upon  which  the  various  points  of  evidence  depci  I 
hrini:  t"  li'.'ht  fads  of  the  \eiy  greatest  importance  in  the  study  of  \\i'- 
jihysiolotry  of  the  In-arlheMt.  One  fact  whiili  stands  out  promiiicii!l> 
is  fliat  tin  iiriiilisl  rlnilliiiiir  fxnvi  r  ri  siili  s  In  Ihi  Ixisnl  imrHini  ol  /  . 
Iii'irt  that  is,  in  «liaf  corresponds,  in  the  more  piimitive  hearts,  to  iIm' 
siniis  vciiosiis, 

Altli.iii'.'h  the  muscle  of  the  entire  heart  possesses  rhvthmie  iiower.  if 
docs  not  do  SCI  to  an  e(pial  dcurce:  in  the  sinus  the  rhythniie  powi'i  ix 
exliaordinarily  developed,  while  in  the  liullius  arteriosus  it  is  scan.  ly 
lecou'iiizaMc.  This  ol(scrv;it ioii  suttiicsts  the  possiliilily  that  the  si-  i^ 
may  dominate  the  he;irt licit  that  it  may  lie  the  "pacemaker"  for  ic 
heart  as  a  whole.  The  most  natural  niclhod  for  dciiionsfraf insr  sin'  :i 
])iissil.ility  woiiiil  he  to  ohservc  the  elTcet  on  the  heartlicat  i>\'  sdiik  h'  I; 
lietwecil  the  sinus  alul  tln'  lest  of  the  heart,  Such  a  liloek  can  lie  in  ■'- 
duced  ill  the  heart  of  cold-lilooded  aiiiniiils  l>\-  local  compression  am  I 
the    \arioiis   junclioiis.      If   a    thread    is    tied    around    'he    siiioauri<'    i'' 

ji tidii,  the  sinus  will   tro  on   liciliii!,'  iiniiiteiriiptedl\-,  hut   the  aim    ■> 

and  vcnfrii-li's  that  is,  the  urcaler  part  of  the  liiarf  helow  the  liual  ■■> 
-  will  cciise  lieatiii'.'.  s.iiiietinies  entirely  (Staniiius'  ligature),  Affi  .1 
while,  however,  the  heart  lu'lou  the  liuatiire  will  usuall.\-  lieirill  to  !■  I. 
Iiuf  at  a  rhvthni  which  is  slower  than,  and  indi'pcudcnt   of,  that   ot     Ur 


Tin;  iiiVMioitMiV  OK  Till:  in  amtiii  \T 


17.- 


'I'lii'  t'\|tri  iinnif  I'liii  In-  Ntill  lifllci'  pi'i  fo.  ini'<|  l(\  nsiiiir  n  wi'iIkc- 
Oiiipcil  clinnp.  ((iiihkcH's  cliiiiiii.  i  If  lliis  is  applied  so  Miiif  tin-  lii-iiit 
r;iM  lio  piiii-lii'il  t'itlicr  tit  till-  sinoinii'iiMiliir  jiiiiclinn  oi'  nl  tlir  tiiii'ii-iil<<- 
V,  iiti'ii'iiliir,    it    will   III!    t'oiitiil   tliat,   as   the   ciMiliac   tissuf    ix   ({"■•luiilly 


I    ^      t'       111  .lit  tiloi  k     |<ti'>liti-t'>l     li>     .i|i|il>iiiK    it.itu|i    al    uv    jLiritlioii.       Till     ihitiip     w.i^     (luliti  in  il 

at    4.      vl'ioni    111  iilmkvr.) 

|iiticlii-il,  till-  purtiiiii  .  iif  lit-ai't  lii'low  Tails  to  l>fiit  as  <|iiifkly  as  that 
iliiiM'  tiif  fiaiiip  tV\K  i'Ji.  This  is  known  as  imrlinl  In <irt  hlnfk,  ami 
Mm-  (If'.'ifc  of  ihf  liloi-k  is  iiiilii'ali  il  li\    till'  iiiiiiii'i'ii-al  fxprt'ssioii  "2  to  I. 

■  '•  to   I.  4  to   I,  t'tc,   iiM'aiiiiit;  that    llic  sinus   is   lifalin<;  fitlicr  twice  as 

■  luicklv  as  the  veiitriele,  or  three  times,  t>i-  foiif  times  as  the  i-ase  may 


vi 


i     If 


i  tk.'.  4.1  Tracili;;  ot  i '.iitr.ii  tioii  ..i  vriiliul..  *:...v\iii«  1  liv  «  Ilr.  I  ..t  llir  I.it.il  .i|>|illt:ilii'ii 
'■ii.it  t.i  till-  aiiruK  at  /,  aii<l  t..  tm-  .ip.-  i>l  tlit-  Viiitiiili-  at  .'.  NmI.-  111. it  tlu  lali'  in 
:i^.  ■!    Ill    till-    f.iriiiir    ia«i-. 

'.     Similar  eonditioiis  of  heart-Mot  k  may  also  he  prmliieeil  liy  eiittinu 
I    eaiiliae  tissue  Jiartly  aeross  at   various  plaees  in   tlie  heart. 
I'liither  evitlenee  that   the  sinus  tloininates   the   heal    in    the   heart   ol" 


f!t'  >   "- 


\l   I 


I 


ITtI 


Tin;  niiii  i.ATiiiv  op  tiii:  lii.ouD 


p      i 


cciIiI-IiIikkIciI  iiiiiiiiaU  is  t'liniislicd  liy  iiliscrviiiy:  tlic  clTt'cts  of  local  heat 
iinr  or  ccnliinjr  of  tin-  vaiious  paits  of  tlw  heart.  In  all  rhytlinii<'all.\ 
;'i-tiiijr  si rui-t nil's  it  is  wcll-kiiowii  that  heat  iiici-casos  the  fate  of  tin 
ih.vlhiii  ami  coM  di-picsscs  it.  If  wo  loi-ally  wanii  the  icjfioii  of  th« 
sinus,  as  liy  holdiiiji  a  hcati'il  wiii-  iii'af  it  the  whole  lieart  will  iiiiiiiedi 
ately  lieat  (luieker;  hut  if  we  locally  heat  the  tip  of  the  veiiti'iele,  ii 
alteialion  of  rhythm  will  he  ohserv  cd  to  occur  (  l-'iir.  4;!i. 

riic  cstaliiishnieiit  of  the  fact  that  the  sinus  doiiliuatos  the  heaitlicr 
that  it  is  the  |)aceiiuikcr  (d"  the  heat — raises  the  qviestiou  as  to  how  1h' 
iMipulse  oriiiiiiati'd  at  this  i)lace  is  transmitted  over  the  rest  of  lh' 
iicait,  and  here  ai;ain  a  neurofjenic  and  a  niyo!;enic  hyjjothesis  have  1" 
he  considered.  Ilcforc  <;oin<:  into  this  (|uestion.  however,  it  will  be  wcM 
lor  ns  to  consider  hrielly  the  nuiuner  of  res))onse  of  cardiac  inuseii 
liiier  to  a  stinndus,  hecause  the  hehavior  <if  cardiac  muscle  uiuler  suci 
conditions  is  considcrahly  ditferent  in  many  rcjrai'tls  from  that  <d'  skcl 
ctai  muscle,  and  it  is  to  these  dilTcrences  that  nuiny  ot  the  ])eculiai 
ali<'rations  in  the  licat  observed  after  interl'eiini,'  with  the  condluMiii- 
structures  lictwccii  the  sinus  ;ind  the  rest  of  the  lieart.  ar'c  to  he  c\ 
phiined. 


The  Physiologic   Characteristics  of  Cardiac  Muscle 

It  is  necessary  to  hrintr  the  heart  into  a  ipiiescent  state  in  order  V 
in\estiy:ate  the  i)ro|ierties  of  its  musculature.  This  is  accomplished  1' 
till'  application  of  the  !<tannins  liirature  lictwcen  the  sinus  and  the  ami 


I'll;.     11       ri"U     I". Ill     -li.iuim;     llu'     |i.-in.iii     ..I     til.      li-l     au.l     -c, I     li,nalur.'<     .it     Sl.ii  i 

'll"t"iii:       '.    :iiiiiil.>;       ,    ^inii-;     ,-.     M-iaii.l.         It     1^    tlu      lii't     liK.itiiu*    whi.Ii    l.riiiRs    tin      li. 

!■.      -I.lll.MllI 

clc-.  I  l''ii_'.  4-1  1.  .\ftcr  tiu-litciiiiiir  the  liu^alure  the  auricles  and  \ ciitrii-i' 
lieconic  oliiesccMt,  .'iiiil  liy  iilisci\in!i  the  ciTects  produced  li\  the  .ipp 
cation  ::f  .'Icctiic  or  other  >tiinuli  we  can  coiripare  the  liehavior  of  '^ 
cardiac  iiiuscle  with  that  of  skeletal  muscle  similarly  stimulated.  'I'l 
ciimpariMin  is  made  lieciusc  of  the  assistance  which  it  nlTers  in  conipi 
hci.iliii".'  the  properties  of  cardiac  iiiiiscle.  As  a  matter  of  fact,  rcc 
in\  -tijraiions  have  slmw  n  that  the  ditVerences  lictwcen  the  two  I\)>cs 
niiisc'e  ,'ire  not    fiindaiiieiital.  since  under  certain  conditions  the  one  iii 


Tin:  ciiYsioi.ddY  nv  Tin:  iii.akthi.at 


177 


mat 


Ic  to  l)''liiivt'   like  the  :illici-.      'I'licv   Jilc   (li'IXMnIfllt    lip""    lllO  ])l'iv 


(ir  iilisclict'  (pf  iiliiistiiliiiisis  liclwccii   llii'   h  .iscic   lilicls. 

Wlit'ii  fU'ctrif  sl'miuH  nl'  varyinu:  stn'ii<;tlis  m .'  iip|)li<'il  to  skclctiil 


III.'    st 


III 


I',  tilt'  (•oiiti'iictiiin  pioiliKM'd  liy  ciirli  sliimiliis  is  |>i'i>p()rti<in!il  ti> 
ifiijjtli  of  tlif  liittfr  (lilt  11  tills  liiis  litcHiiic  of  siicii  a  strcnsith  that 
axiiiial  response  is  elieileil.  In  cardiae  iiiiisele,  on  the  other  haiiil. 
,,  ciitii-ely  (lilTeielit  result  is  olitained,  for  the  weakest  stiinllhis,  if  it 
|.ini|iices  any  resiionse  at  all.  inodiices  one  that  is  iiiaxinial;  that  is.  the 
K.  iiilit  of  eontraetion  is  the  same  as  it  would  have  heen  had  a  miieh 
^t|;,|ltrer  stiiiuilus  been  applied.  Kxi)ressiiiu'  this  result  in  «eneral  terms. 
>  r  iiiav  sav  that  in  cardial!  4iiusele  ii  iiiiiiiiiKil  stiiiuihi.f  iinidiirrs  a  itin.n- 


.1.  — Skeletal    Muscle 


/).      l.'iir,liiU-     MumU 

lltliMs    of    >liMnili    "I     iiur.a«inK    -ti.imili    ..u    Ail.l.il     aii.l    lanlMi-    niii-ili'    I"    illu-lial. 
llu'    "all    111-    li'.lhiim"     |iiiiKV|.lo     ill     tin-     laid  r.        ll'i..ili     /'id,  (T,  j/     /''iv.tiii,'...;: .  I 

'/)((/,  whereas  in  skeletal,  the  elTeet.  as  measured  l>y  the  hei<rht  of 
ratlioii,  is  proportitinal  to  the  iiiteiisit>  of  stimulation.  This  is  soiiie- 
-  known  as  the  "all  or  niithiiifr  ))lieiiomenon  "  (  l-'iir.  4.)^. 
If  iiiaximal  stiiiiiili  are  applied  siieeessivel.\  ami  at  slitir*  intervals 
line  to  .skeletal  muscle,  a  siiu'litlx  liiirher  resjionse  results  from  each 
'•rdiiii;  stimulus,  until  alioul  ten  stiiiiiili  have  hern  a|)plietl.  after 
ill  t'or  .some  coiisitit  lalile  time  the  same  heiirht  of  eontraetion  follows 

i    ^liiiniliis.      If  eael ntraction    is   reeordetl.   it    will   lie  seen   that   the 

'   fi'u  eoiifractioiis  jrixe  a  stain'ase  elfect  :  that  is.  if  a  horizontal  hue  is 
-Ml  from  the  top  of  each   contraction   to  the  next  one,   the  elVect  of  a 


-•    f 


*i         1 
n         { 

I   i| 

1  Mi 

If 

5     III 


Os  jJ"^' 


ITS 


Tin:  ciucn.ATioN  of  tiii;  iii.oon 


f  ^ 


stuii'ciiM'  witli  ^'railiiiilly  iliiiiiiiisliiiiir  steps  will  Ijc  pi'odiiccd.  If  we  rt'pt:  ! 
this  olisiTvatiiiii  witli  i-aniiac  imiscic,  we  sliall  find  tliat  tlic  stairca  ■ 
plii'iiiinii'iKHi  oi'  //•(/(/«.  as  it  is  calltMl,  is  very  pnniiniiiccil ;  and  iiiorcovi!-. 
ill  (ilicdii'iicc  to  till'  all  111'  iiiptirniir  |)riiU'i]ili'.  tlic  ti'i'iipc  is  olitaiiicil  in 
canliai'  niiisclc  \vliatt'\  ci'  may  I»'  the  I't'iativt'  sti'cliutlis  of  the  slinm  i 
applii'il  to  the  lit-ai't,  |Ho\iil('il  always  tliat  all  of  tliciii  ai'c  ('lT<'cti\': 
wln'icas  ill  till'  case  of  skeletal  iiiiiseli'  it  ran  lie  (Icinonst rated  only  jir  > 
\ideil  till'  stiiniili  arc  of  cipial  slreiii;tli  (  l''iir.  4(>i. 

:>.    If  all  elVertive  stililllllls  is  applied  to  a  skeletal   lllllseie  while  ill   priier-.^ 


S'Kfltlal    muscle 


I'artliiit"    inuscU* 

I'iK.    4'.,      'Ih.    ill.ttv    nf   sii.t.-vivc    ^iiiiuili    ni,    sl.Utitl    ,-iii<i    f.ii.liac    iiHistl,    to    nIh-u    llu'    |>t..iiir. 
ill     III!     -laii.a-r    |.lii HI, .11.    c.r    »i.//..    in    llu     l.itlir.       ll'n.m    T.    Ci.     Ilrciclu.) 


I 


of  eoiitraetioii.  as  in  response  to  a  preeediii'/  stiinnlns.  the  seeoiid  stiiiiiil  . 
priiioiii:s  the  eoiitraetioii  pi'iiiluied  liy  the  first  one.     If.  however,  the  sef, 
stililllllls  is  applied  diiriii<.'  the  latent  period"  of  the  first  one.  it  will  have  :  ■ 
etTect      that   is,  the  iiinselc  duiiiiji-  this  period  is  rcfrai'tory. t      Fioiii  tli 
results  it  fiijjiiws  that  stimuli  siieeei'dini:  faeli  other  diiriii'r  the  contrael 
period  will,  ill  the  ease  nf  .s7,(  /<  /<(/  iiiiisele.  eaiise  a  eoiif  iniions  eoiitraetioii.     I 
tetanus,  as  it   is  ealled,  iieeanse  the  eoiitraetioii   prndiieed   by  eaeli  slin    i 
Ills  will  add   itself  to  that   of  its  |ireileeessor  liefoie  any  trace  of  rel.   . 
atioii   has  set    ill.      If,  liuwever,  the  second  stiiiinlils  is  applied  so  lati 
the  contr.-ietioii  jierind  iif  the  lirst  that  time  is  not  availalile  fur  the  lai 

''•.•      "»••!«  Ill      J.ill.Mi"      i~     iHi.ini      (in      |'irt...|     .111(1      111,       Mil. UK  lit      nl'      .1]  i|  >I  l(  .ll  j  i  HI      t,i     a     Stililllllls     <j 
Uhilh     Till    ttfl.t    nf     tll.lt     ^tPll'lll!^     i~    i.lis,  I  Vf.l. 

tl'.y   "r<iiail..ry    |.,!in,|'   ,,    Mit.iiit    tin    liiiu-   |.,l|.,Miri:    tlic  .i].|.lii  ati.iii    ..f  a   sliinuln~   .lliriiii;   « !: 
w.r,i,l    siiiiiiiliis  .l,A<l,.|.»  1,-s  lli.iii   Its   full  <lt.il   ..r  11..  .iri.l    .It   all. 


Tin:  I'liYsioi.odv  iiF  Tin;  in;AKTi!i:AT 


170 


IMiiod  t(t  fXiMMid  itself,  llion  (il)vi(iusly  a  slifrlit  rclaxiitioii  will  liiwc  oc- 
iiiirc'l  hcfiiro  the  olTcct  oF  flit'  second  stiiiiulus  develops  itself,  and  tot- 
jiMiis  will  he  incoi.ii)lete.  These  faets  will  lie  evident  from  the  aceoni- 
liiiiiyiii<f  traeinfrs  (Fifj.  47). 


"S 


r. 


Sk.ltt.ll     HIUmIi- 


Staiiiiiiis'   lu.irt 


liy.  47  Tile  iffirts  t»f  Slier*  s^ivi'  stiniiiii  .'iioi  >'!  t(  lani/ini;  ^tininli  .-n  skri*  tal  ilnKcIo  nini 
T  l.u  iiuis.!.'.  Tin-  •.mall  v<t':.:iI  iii.irk*  ^ll.■w  mIuii  lli.-  ^liiuiili  win-  iiilr.i.lu,  i-d.  (l'iiiii|,il.  .1 
"'.    ll.iui.i;,     |.iii.lMu-.l     l.y     T.     C.     Ilr..,ii.      nil.;      i.<.ii,ir.l     Kill.  I 

111  the  ease  of  cardinc  muscle  the  aliovc  dcscrilicd  |>i'o|i('i'ties  are  (|uite 
M'Tciit.  for  llif  n fnivtori/  pluixi  (  rh  mix  Ihrinuihuiit  tin  irlmh  fiiriiiil  tif 
■'tiriiiliiiii ;  that  is,  a  second  stimulus  applied  diiiinir  the  coiitraetion 
'I'liuced  liy  a   pie\  ions  stimulus  has  no  elTeet   whatsoever;   it   does  not 


h 


t 


18U 


■IIIK    tlKII  I. Alius    (IK     I'lli:    lll.(M(ll 


liavc  line  until  llic  iniisrlr  li;is  t'c'irlicd  llic  I'lill  cNtt'iil  (iC  its  fiiiiti'a<-tii>ii 
aiul  is  aliDiit  to  relax.  Sim-c  a  latent  period  must  supervene  upon  the 
np])lieation  of  this  seeoiid  stimulus,  it  t'ollo\vs  that  no  eoniplete  fusion  oi 
tile  eontiaetioiis  is  |)ossilile.  Coiiiplete  tetanus  tlierefore,  does  init  oeeiii 
ill  eardiae  iiiilsele.  liowcvei-  frei|iieiitly  the  stimuli  may  lie  ap|)lied  t  l'"i;r 
47  I. 

Tiie  refractory  ])liase  is  a  ]nopert.\  of  extreme  inii)oitaiM'e  in  inider 
standing  many  <if  tlii'  peeiiliar  irrejiularities  olisei'ved  in  eardiae  aetioii. 
If  «e  oltserve  the  effect  of  stinuili  applied  at  vai'viiiff  periods  after  th  ■ 


'i  i 


I'm.  -IS.  -Myonraiii^  of  frn^'s  vcnti  iiU-.  ^tmwinK  rtftct  of  excitation  !)y  break  indue ti'> 
^liink".  at  various  nioim-nts  ^^•  (he  cartliai'  cytle.  Tlif  line  O  iiulicates  tin-  Cfunmenccnient  ■ 
.ill  ilu'  ht-at-i  (InriiiR  wliicli  tht-  ^Iiock  is  Pcut  in.  It  will  lie  noteil  that  in  ;,  ^  and  ».  the  ht-.r  ■ 
is  rtfr.iitorv  to  tti<-  -tiinnlii'*.  The  si^naK  iii'lii.iic  ilit-  nioiiu'iits  at  which  ilie  stiimili  were  .i 
I'lii-d.  r'ruin  4  to  V  tlu  heart  reacts  hy  an  ;\trasys'nh-.  after  a  delay,  which  is  proKfi'^^sively  li- 
the later  in  diastole  the  siir.udii'i  enters,  as  shown  hy  the  sections  shaded  oliliiniely  to  make  tli-  ■ 
iiiiiie  I  on!*iiiciions.  The  evtra^ystoK  ■;  tiicriase  in  height  from  4  to  .?,  each  heinR  followed 
,1    Cfiiiinii-iatory     iiao'-e.       i  l"ri<m     (,im  iaiiiS    Ihnnan     I'hysiology.) 

tt'i'iniiiatinn  of  the  ict'iartory  i>lias('  of  a  inoxiotis  stinuilus,  ue  shall  lii" 
that  the  lu'ijiht  o*'  'he  extra  (•(Hitractioii  is  <iir(M'tl\  pr<tintrtitnial  to  tl 
tiiiH'  at'lt  }•  tln'  i'IhI  ot"  \\iv  tt'l'rat'tniy  period  at  which  it  is  ai)pli«Ml.  If 
stinmliis  is  applied  at  the  vei'.v  he^riimiii*^'  of  diastole,  the  extra  etnilia' 
tioM  will  he  small,  w  liereas  if  it  is  applieil  at  the  end  fd'  diastole,  tl 
extra  eoiiti-a4'titni  will  he  at  least  as  hiyh  as  that  of  the  ])feeediii^.  I 
may  he  liiirher  heeaiise  of  the  ti'ep[>e. 


"i 


Tin:  I'HYsioi.odY  op  Tin:  iikahtiikat 


181 


These  obscrviitidiis  onnlilc  ns  to  intci|)it't  the  results  oliliiiiifd  l>y  c/*- 
liliiliifi  rlccir'w  sliorks  (<.rtni  stiiiiiili)  to  thr  hnitiiii/  linirf  (lui'ins;  diffofpiit 
[ihiiscs  of  systolt'  and  diiistnlr.  Diii'iiij;  systole,  the  iimscle  heiiij;  rcfnie- 
tniy,  no  efTeet  is  ptodueed  Im  the  extra  stiiniihis.  Imt  duriii-f  diastole 
lAtia   systoles  wliieh   are   i)roi;ressively   more   ])roiiouneed   the   later  in 


tin 


rnr,  fii 


.f 


eh  sti 


iih 


Thes 


iliastole  ttiey  oeenr,  loMo\ 
Niilts  aie  so  far  exactly  like  those  obtained  with  a  ((iiieseent  heart.  But 
iiiiplliei-  ]>henoiiienon  now  heeonies  f\  ident  ;  namely,  that  following  each 
.  \(ra  systole  then-  is  a  eom])ensatory  pause  in  the  aetiou  of  the  heart, 

■  if  such  duration  that,  when  the  lu-xt  natural  i)eat  occurs,  it  does  so 
|ii,icticaliy  at  the  same  time  as  it  would  have  occurred  had  no  artitieinl 
viiiiiiiius  licen  applied.  This  will  lie  apparent  from  the  followin;^  dia- 
'..iiaiii  t  {''iii.  48). 

It  should  he  noted  that  the  refractory  period  is  ^'reatly  dinnnishcd  hy 
r.ii-injr  the  tcmixTaturc  of  the  heart.  Indeed,  under-  these  conditions 
.iii'i  \\ith  stronir  stimulation  it  lua.v  lie  jxissiMe  to  i>roduce  an  almost 
'Miiiplcte  tetanus. 

The   iin])oi'taiu-e   of  hnowiuu'   the  aliove   facts   is   that    \\v   aie   thereb.v 

■  ii.ilited  to  eN|)lain  the  peculiar  niainier  in  which  the  \entricie  responds 
t.i  stimuli  transmitted  to  it  from  the  sinus  and  the  auricle.  The  museu- 
inlurc  of  the  auricle  and  xcntricle  of  the  mammalian  heart  is  not  one 
■■'iiitinuous  sheet,  luit  is  sejiarated  liy  a  sjiace  at  the  aurieuloventrieidar 
iniietion.  across  which,  in  specially  oru'anized  structures,  the  heat  of  the 
I'lricle  is  transmitted  to  the  ventricle,  .^ometiiues  the  stimuli  are  so 
iVcipicut  that  the  ventricular  muscle  is  uinihle  to  resjiond  to  each  stimu- 
lus transmitted  to  it,  with  the  residt  that  marked  irres:ularities  in  con- 
'raction  occur  (see  paye  L'sot.  In  this  way  certain  of  the  cardiac  irreiru- 
'arities  ohserved  in  nuin  can  lie  explained.  Thus,  the  so-called  iiiilxiix 
''I'lmiiniis  is  due  to  every  second  heat  lieiii-x  an  extra  systole.  This  second 
'•at  is  therefore  generally  weaker  than  the  preceding.'  and  succeedin;;  nor- 
■lial  lieats,  and  it  is  almost  always  followed  liy  a  compensatory  jiause.  When 
;lic  intervals  separating  the  lieats  are  of  uniform  leuirth,  although  every 
-cond  heat  is  diminished  in  size,  the  pulse  is  ternu'd  iiiihiis  altcnians. 


r 


Lj-  Vi 


!'  » 


(IIAITKK  XXI 

TIIK  I'I1VS|(»I,()(;Y  OKTIIK  llllAUTr.KAT  (CiiiitM) 

THE  ORIGIN  AND  PROPAGATION  OF  THE  BEAT  IN  THE 
MAMMALIAN  HEART 

As     llMS     llCIII     nIiDWII     ill     lllC     prfl-Cllillir     clliiptlT.     tluTC     is     Ml)     llolllll      lllilt 

ill  till'  cdhl  IiIikkIiiI  liKtrl  till'  liiMt  ori'.'iiiiili's  at  tlic  siims  vciicisns,  wIicium' 
it  spicads  til  llif  list  of  tlic  liciirt.  \'ci>  stnnij;  (•\iilriii'c  lifis  also  ln'cii 
pii'si'iili'il  to  imlicali-  lliat  tin-  lifatiiii;  |>o\\(r  is  iiilwi  i-iit  in  llic  inuscli' 
filiiT  ifsrlf  ami  iiiili'|H'ii(lfiit  of  iiiMvoiis  stnu-tiifc.  Tliis  woiiM  sn^'-ri'st  tin' 
I'lirlliiT  iHissiliilit.v  lluii  tlic  structuii's  lliroutrli  wliicli  tlir  ln'at  is  iiropa 
irati'il  ai-f  till'  iiiiisi-lf  fillers  and  not  tlii'  nerve  liln'i's  in  other  \\ofils, 
tlial  tlie  |iro|iauatioii  oi'  tlie  lii'artlieat,  like  its  oriyinatimi.  is  ni\o<tenie 
I'atlier  than  neiiio<ieiiie,  l>ireet  prool'  ol'  tliis  livjiotliesis  is  readily  fur 
iiislieil  liy  imiiieroiis  expeiinicnts,  aiiionu'  w  liieli  may  lie  nieiifioiied  mak 
ill"  iiilerditritatiiiLr  cuts  across  tlic  iieart.  or  excisiiis;  a  rililmii  of  vcii 
tiiciilar  muscle  liy  an  incision  simiilatini.'  tlic  walls  of  Troy.  Tii  liotli 
tlicse  cases  tlic  Iicat  will  lie  found  to  travel  fioiti  one  end  of  the  imiscular 
hand  to  the  other,  althoinih  it  is  eviilcnt  that  all  the  nerves  procccdiiiL' 
from  hase  to  apex  of  the  heart  must  have  lieeii  severed.  Of  course  this 
e\  ideiice  is  Hot  irret  iitahlc,  for  it  mifiht  lie  aifiued  that  there  are  nerv- 
ous stiiictiiics  disposed  ill  the  form  of  a  plexus  continuously  all  over  the 
heart,  and  that  some  liranclics  of  the  iilexus  remain  uncut  in  the  aliox.' 
experiments.  It  is  only  in  the  heart  of  Limnlus  that  uiidouhtcd  evideiicr 
exists  that  the  heat  is  transmitted  liy  nei\es,  liut  as  w  c  have  seen,  this 
heart    ill   all    its   pro|ieities   is   jiiolialily   the    proxcrliial    cxceiition    which 

plo\cs   the    rule.      The  halaiic f  evidence   stands   in    favor  of  the   vie\' 

that  the  propauation  of  the  heal  o\er  the  cold  lilnoded  heart  is  myoj^'cni' 
and  not  iieiini'j-''iiic. 

CONDUCTING  TISSUE  IN  MAMMALIAN  HEART 

When  x\  e  attempt  to  iii\ cstiuate  the  prolilems  of  the  oriiiin  and  prop," 
■nation  <if  the  lie.it    ill  the  ir(iniilil(iii(l(  d  liiarl.  maii,\'  experimental   ditV 

cnilies  111  collide  lac-  IIS.  in  o\  el  com  iie^  llic>e.  the  ii  I  >l  thili'i  We  liili-' 
do  is  to  estalilish  the  structural  relationship  lietween  cohi-liloodcd  an- 
x\  arm-lilooih'd    hearts.      In   the   emlir\o   of   hoth   classes   of  animals   tli' 


isj 


Tin;  i'iiYsi(ii.(MJY  OK  TiiK  iii:.\hthi:at 


183 


it    iii'iscs   fis   tlu'   SD-ciilltMl    <-ar(liiii-   tiilic.      As   (Icvclupiiii-iit    procct'ds, 


(li^citit'iila  >{i()\v  (Hit  t'lipiii  the  walls  of  tliis  tulif  to  form  lli 


<■  auni'lcs  anil 

\.  litiii'li's.  Ill  the  comparalivfly  simple  licart  of  tlic  tiiitlc  tlicsc  dispo- 
xiiiiiiis  of  the  aiiriclis  and  \ fnti'iclcs  in  ?'"'!.itionsliip  to  the  cardiac  tiil)c 
an  iiKirc  or  less  evident  even  in  tlie  fully  developed  lieart,  |)articiilaHy 
ill  the  ease  of  the  auricles  (  l-'ii:.  4!l  i :  liut  in  tlie  licart  of  tlie  liijrlier 
iiiaiiiiiialia  it  is  impossihle  liy  snperticial  cxaiiiination  alone  to  show  any 
i.  hi.iins  of  the  primitive  cardiac  tnltc.  .More  carefid  anatomie  iii\ cstifia- 
tM.ii<  diirinu:  recent  years  have,  however,  shown  that  it  exists  in  the  fcniii 
•  ii  ciitaiii  (Iclinite  structures  composed  of  tissue  liistoloy:ii'ally  <piite  dif- 
ii  rriit  from  that  of  the  icst  of  the  iieait,  and  disposed  in  such  a  manner 


.!-  Would  indicate  not  only  that  it  is  derived  from  the  |)riiiiilive  eardiiic 
:  lie.  Iiiit  also  that  it  is  the  main  pathway  alon;;  whicli  the  heat  is 
'.'  iiiMiiitted. 

I  ills  priiiiilive  cardiac  tissue  is  much    lietter  dc\cli'i)ed   in   certain    re- 

■  "Ms  tliaii  in  others,  the  lirsl    p(n-tion  of  it   to  he  disco\cred   lieinu;  that 

'  i"wii  as  the  (itirici(l(ir(  iitriniliir  iimli,  or  tiie  node  of  Slanley  Kent*  I  Fi^'s. 
'  'iiid  ■)!  1.     This  structure  is  found  at   :lie  liase  of  the  iiiteranriciilMr  sep- 

■  :ii  111!  tile  ri^.'ht  side  and  near  its  po>terioi'  iiiarjrin.  It  exists  as  a  Meetion 
"'■  peculiar  small  primitive  cells  and  fihers.  and  is  continued  downward  as 
'I  hi'iiilh  of  the  same  iieeuliar  tissue  to  the  interventricular  sepfuni, 
vicic.  near  the   iinioii   of  the   jiosterior  and   median    Haps   of  the   aortic 

■|  .'    aiMuviry   ..(   llLi>   II,., I,    is  of. 11   trr,jm,,uv|.v   atlriimt.,1   I,,   Hi-.  mu\  ..ill., I    ,ilur   Iji-   tmiii.'. 


J 


L..J> 


-r^ 


Ifit      t 


t   I 


184 


TIIK   ni«l  I.ATlnN    OK    rili;    lll.iioii 


valvf,  it  liiriii'i'iili's  .so  jis  III  M'lul  II  lii-iiiicli  down  cacli  side  of  the  scptni' 
iiiini<>iliatfl\'  Ix'low  tlic  ciiilofiiriliiiin.  Ividi  iiiiiin  liraiicli.  as  it  procp(>il 
downward  on  the  Ncptuiii.  divides  up  into  an  intiicatc  system  of  sinalli  - 
liraiielies,  wliieli  l)eeonie  retlerted  ovei-  tlic  inner  surface  of  tile  ventrieli  - 
>\  liere  tlieir  existence  lias   liecn    known    lor   mmiic   time   as   the   so-eallc  i 


Fir.     'If        Ih-'i>t)iin    ..t    'm.ih    tn      li.'U    .uiiii  nl"VMiIi  it  nl.n     I'limlU     iKiltii';    .*.    tlic    l.tKiii 
ilic  1. nil. II,.   kiM.vMi  .,-  il„.   .\ A    i,.„l,  ;    J,  tl„.  1,,:,„|1,.   ,li>i,liiiK    Mi.   t«..   Iii.n»li,»;   4,   llif   l.raii. 

illMB     "II      t'lr      riKht     ^l.if      i.l      thr     ■■!?■   t-v-,  .;iru-,lla|-      *i;i('llll.         (  l"rnm      ll.iv^lir^      /'/!\..f.i'   'r/\,  I 


ft 


in:      >1.      I'!int.inr.-t|.li    Ml     111, I, 111    Mt     liif    aiiriciiluw  pilri(  iil.tr    Inimllr    .tn.l     il^     t.Tinifirali.'ii-. 
-Ini.lril    fn.ni    .li^^rc  tioii-    .1    llir    In  m!  I    (  Mi^s    l)r    Willi.      All    of    thr    lir.imlir,    in    tin-    Irft    vii- 
,iri'    not    ill,  hulcl.      I  l-'r,.n:    l|,.w,ll  i 

I'urkinji  fihi  IS.  Tln'  tiliei-s  iiltiniiitely  end  in  close  as.sociatioii  with  ' 
pajiiliary  muscles.  Tin-  node  ami  main  luiiidle  ami  tlip  two  liram'l 
lioforc  tiioy  iiave  licirmi  i,i  divide  aie  siii  ruuiidrd  by  fihrous  ti.ssuo,  .. 

tliey  seem  to  have  a  lilieial  lil 1  siippl\.     It  is  of  interest  that  they  (■'■ 

tain  a  hij/li  iicrceiita^rc  of  u'lvcoyen.     In  llie  human  heart   the  aurien' 


TIIK   I'HyHUtl.lKlV    (IK    Till;    lIlAltrillAT 


isn 


[III 

'Cil 

II. 
■I. 
Ill' 


m 


M 


'  nti'iciiltii'  iiiiijc  iiikI  iiuiullc  iiit'iiHiirc  aliiiul   l'>  nil 
J  Mini,  in  w  idtli. 


I<iit;1li  aiut  ahoiit 


Till'  lest  of  tlif  lissnc  lictwc  II  tlic  iiiii'ii-li's  mill   vciitricb 


fill 


I'l  iiiifiirf,  .•iltliiiiii.'li  oflicr  ciiiiiH'rtiiHis  likf  tllll^('  of  lin'  aiiriciilovonfricular 
!  iiiiiilf  liavc        •!  (Ii'si'ribod  liy  Kent.     Oiir  of  tlii-isc.  i-allcd   IIk    rii/ht  laf- 

■  r,tl  iitniiii'lion.  niiis  lictw cell  tlir  rijrlit  iiiiiirlr  .•mil  tlic  external  wall  of 
I  ill'  riu'lit  \eiitriele. 

Aniillier,  liiit  niiieli  smaller,  iiimss  of  similar  einlitvonie  eardiiic  tissue 
!  ;is  nmre  recently  lieeii  diseoM'red  liy  Keitli  and  KlaeU  in  the  parts  of 
till'  auricle  wliicti  correspond  anatoinieally  to  the  sinns  xcnosiis  of  tlie 
iii'iirt  of  ciilil-liliioded  animals  tliat  is,  in  tlie  area  Ixinu'  lietween  tlie 
iipi'iiin!.'s  of  the  vcine  cinie  and  amnnd  the  coronary  siiiiis.  To  lie  more 
i\|'licil,  tliis  tissue  lies  "in  the  sulcus  terniiiialis  just  lielow  the  fork 
iHinieil  liy  the  .junction  of  the  upjicr  surface  of  the  auricular  appendix 
•itii  the  sujieriiir  \ena  i'a\a."  This  .ilnotiiirlriiliir  noili,  as  it  is  called. 
I-.  more  or  less  cluli-sha|)cd,  tlie  Mliiit   end   of  the  ciuli   licili!;  alxix-e,  as 

^Imivmi   in  the  a inipaiiyin<„'  fiirure  !  Kii;-.   .'>2i.      It    is   iiii|)ortant   tu   nnte 

iluii  tlii'i'c  is  no  direct  connection  visible  lietween  llie  sinoauricular  and 
.niriciiloventricula?'  nodes  ( i-'iir.  ")•'•'. 

.\niither  anatomic  fact  seen  jilso  in  the  acciimpaii\  in;r  fipiire.  cnnceriis 
'111'  ilispositioii  of  the  muscular  lilicrs  of  llie  auricU'.  Tiiese  radiate  in 
iiiiidies  in  a  )ieeuliar  fan-shaped  manner  from  a  iioiiit  wliicli  lies  im- 
111  iji.'iti'lv  lielii\\  the  sinoanricular  node  to  all  parts  of  the  superficies  of 
I  III'  liirht  auricle.  This  point  lias  lierii  called  the  conn  iitratloii  point. 
.\\  the  "rininatioii  of  the  vcire  cava-,  the  muscle  fillers  are  arraiiired  more 
"I   los  circularly. 

IIa\iii<;  hocoiiie  familiar  uitli  tlie  disposition  in  the  mairmaliaii  heart 

'  !'  the  primitive  cardiac  tissue,  aloni;  which   in  tin-  heart   of  the  lower 

.iniiiials  we  know  that   tlw  lieartlicat   s|),'eads,  w c  ma.v   now    proeeid  to 

'  \aiiiine  the  evidence  sliowiii'.;  that  this  tissue  is  also  res])onsil>le  for  the 

"rr.'iiiiitioii  and  propasratioii  of  tlie  heat  in  the  heart  of  mammals.     With 

■'  -.iiiril  to  tlie  oriirin  of  tlic  heat  in  a  normally  heatiii!:  niaiiimalian  heart, 

'  1^  of  course  iiiipossilile  to  tell  where  this  takes  place.     Tf  tlie  heart  is 

'  Ai'iscd,  however,  it  will  eontiniic  to  lieai    i.  •  a  few  mmnents.  and  as  it^* 

'ics  it  \\  ill  he  ohserved  that  the  iioucr  of  i-oiitraction  remains  in  the  ;i^ 

'  ■  ular  rciiidii,  and  /niitit  iiliirli/  at  the  hases  of  the  veiue  cava',  for  a  con; 

ii  r-ihlr  time  after  the  ventricles  lia\e  ceased  to  heat.     Tiiis  jiart — the 

■mnnti  woririis     is  situated  ill   most   hearts  snmewhat   lower  than   the 

i'i;iiirioular  node.     That   it   is  the  last   part   of  the  heart  to  cease  eon 

■  ■'lih).'  dues  not    necessarily   mean   tiiat    it    is   the   jiart   of  tlie   heart    in 
'  I' h  the  beat   ordinarily  (uijrinates;   it   means  simply   that   this  is  the 

!  nt  uf  the  auricle  in  which  the  imwcr  of  contraction  remains  for  the 


•1    f 


1H6 


Tin:   I  lUCI  I.ATIIIN    OK   Tin:    iii,«mii» 


IdliKot  liiiic  iiflcr  ilciilli.  .\Ii1m(1Ii,'1i  lln'  iiIimtn  iitioM  ilnfs  imt  fimliU-  ii^ 
to  (li'tcriiiiiii'  tXiK'lly  \\\\i-\v  llir  litiiitliriit  (iriKiiiiitrs,  \  el  it  makes  il 
very  pinlttililt'  lluil  llii^  is  soiiicw  iicic  in  tin'  nurii-lt's;  a  coiiclusinii  uliicii 
is  lior'iit'  out  liy  many  otlirr  iiicccs  ui'  cv  iilcMcc.  siicii  as  tliosc  iii)taiiitM|  liy 


I'll.-.     ■'        Hi.itiain    "I     .,11    .11111. Ir     -ll.nliiil!    111.-    an. main!!  lit    "I     llir    iiiu->  I.     ImilN;     Hi.     i-..i„  ,  i 
iriii.ii    |..iiii    11. /'i:    iiiu    111.    ..uiluif   ..1    ihi-   S.J.    ii...ii     i.s,l.\.i,      Tlu-   ih.iwi.iiii    in    In   -lal..   .ii. 

illii-ti.iii-   l\    III.    .11.!.-  !   ...111...  tiiii!  .I.iiii.l   liii.>  till-  Mnth...|   ..f  li.iliim  .ill  l.y   i.air.il  i..iii,i. 

.ill  I    III.     -111.-.  11111  III    .'111  III. .11'  II.       tj-r.-ii;    Thmii.t-    I..\m-1 


^;f 


..■Stnoturlcular  node 


.  Auriojltventrlcular  node 
.Aurkuloventticulir  bundle 

Fight  a  left  ventrlculir 
bundlei 


LJL-Muxuii  pjpii'lares 


Mi;,    -.1.      liiaiTini    t-.    -tn.u    ilir    !■.  n.  r.il    i.Tiiil  .  ali...i«    ..I    Hi.     i  ..ii.lii.  liiii;    li--iii-    in    Itir    lir.iil 
111.-    iii.iiiiiii.il.      Il    «ill    111     ..li-.m.l    thai    III.  1.     1-    ""I'l     1.1    llii>    li--ni     l.ilw.iii    tin     Mnn.iiiruiilii     .. 
.'iiiri.iilnv.  iitri.  ii'ar    ii-.iii  - 

tilt'  stiitly  lit'  |ii>lys|i|iyy;Miii<.'iaiii.s  i  |i.i;.'c  27:>).  nt'  ilol  riN-ariiiii;;i'ams  ( |i:i 
'Jfii'ii.  illi.i  ill'  iihscrv.-itinii.s  nil  llic  li.-.itl  liufiti;;  lit-al't  lilnt-k  (j>at:f  -T-' 
Our  iiiiiiiifiii  tiicrt't'iiic  iiarniws  itsflf  iIdwii  tii  ilfti'rminiii};  tlu'  cx.i 
jxiiiil  III"  till-  riirlit  aiiriflf  at  uliii'h  tiif  ln'at  tniKiiiatcs. 


Till:  I'livsKiiiHiY  (IK  Tin;  iikaktiikat 


SITE  or  ORIGIN  OF  THE  BEAT 


187 


The  «(irkiiiir  li\  potlicsiN  rrom  «liifli  we  niav  pniccrtl  to  attiick  lliin 
I'l^lili'iii  is  that  the  beat  uriKiiiatcH  in  tin*  siiioaiiriculai-  iioilt',  and  to 
|Mi!  this  to  till'  tt'st,  vaiious  iiii'tii<Mls  lia\r  liccii  ciiipioyi'il:  (1)  Wanning 
III  riMilini;  or  iiijiiiiii!;  tlio  iioilr  ami  iintini;  tlic  clVn-t  on  the  iD'ai'tiicat. 

Si|ili  |)i cdiin's  ifccatly  afTi'ct  tlu'  rate  of  tlic  lifai'tiit'at,  wlicrcas  they 

|iiiiiiiicc  iKi  chaiiuc  when  applii'tl  tn  othrr  iiarls  of  the  ht-art.  CJ)  Dc- 
til  iiiiiiatinii  of  tJir  i-niiiparativi'  rliytiiinic  iiowcf  of  strips  cut  out  from 
iliiTi  itiit  rt'tjions  of  tin-  aiiriciiliir  walls.  It  is  jrrcatcst  in  thoso  taken 
iiniii  llif  rcjiii'ii  of  till-  noilf.  (ill  Dctcriiiination  liy  tin-  use  of  i^alvan- 
(iiiicliic  ciiivcs  of  tlir  relation  of  the  iioile  to  the  seat  of  ori'_'in  of  cardiae 
iiii|iiiUe.  \'>\  all  these  inethoils  the  results  indicate  clearly  that  the  heat 
iiii'.'iiiiitcs  ill  the  siiioaiiricnlar  node,  hut  on  account  of  the  creat  iiii- 
piiitaiice  ill  eoniieetion  with  the  interpretation  of  elect i-ocardiofjrains  in 
iiKiii.  it  is  particularly  with  the  result  of  the  third  (.'loup  of  experiments 
Il,,il   «e  «ill  coiiceiii   ourselves  here. 

Evidence  Furnished  by  Studying  the  Current  of  Action  Which 
Accompanies  the  Heartbeat 

To  start  with,  it  is  essential  that  we  niake  oursehcs  familiar  with 
//"  /'/•(ii('i///«\  of  till  tiiitliDtlx  ein|iloyed.  These  principles  are  hrielly  as 
i.illiiws:  When  a  wave  of  contraction  passes  aloiiir  a  iiiuscle,  it  is  ini- 
inriliiitely  ]>reeeded  hy  a  chaime  in  electrical  potential,  which  can  he 
lii  licted  liy  means  of  a  tralvaiioinetcr  connected  with  th<'  muscle  through 
>'.illiil  iionpolari/.iilile  electrodes.  The  f;al\anometer  employed  must 
111  •■\tieiiicly  sensitive,  ami  must  not  vilirale  after  the  current  has  ceased 
ti.  |i.iss.  The  form  ireiierally  in  use  tod;iy  is  known  as  llu  slriii;/  tinli'ii- 
,:,.„ii  I,  r  iif  Kiiilliomi.  It  dilVers  from  the  (.'alvaiiomcter  ordinarily  eiii- 
i'''V'!  ill  ))liysical  lahoratories  in  that  the  current  instead  of  passing 
ti  I'U'.'li  a  coil  of  wire  siirroumliny;  a  iiiatrnctic  neeille.  passes  throuvjli  a 
■^J  •li/.ed  (pM't/  thread  suspendeil  in  the  stroiiir  ruau'iietie  (ieUl  which 
i\'^t>  hetwei  II  the  two  oi)posiuir  ])oles  of  a  horseshoe  elect  roiiiajjnet. 
T  ■  stiiiii;  is  thus  surrounded  on  all  sides  hy  iiinumi'ialile  lines  of  force 
'  '1111111^'  hetween  the  two  poles  of  tlie  mav'iiet.  When  a  eiiirelit.  Iiow- 
'  r  siii.ill.  |)asses  alolij^  the  strintr,  it  will  jienerate  lines  of  t'orce  of  its 
' •■  I  iiiid  these  liy  rcactiii!.'  witii  the  statioiuiry  lines  of  force  of  the  field 
'.  :  I'.iiise  the  slrinj;  to  move.  The  strin;:  is  placed  in  the  pathway  of  a 
^'  :  '-'  heaiii  of  lij;ht.  and  its  shiidow,  after  iiein^  mairnilied  hy  lenses, 
:  '".iet'tcd  ini  a  moving'  plioio^rapiiic  phile  or  paper  arran^'cd  in  a 
'■!'■  iiolder.  The  iionpoh.rizahle  electrodes  relVrred  to  are  employed 
I!    ■   iii'c  of  ordinarv  electrotles  in  order  to  ohviate  the  lieneration  of  clec- 


ll      I 


1H8 


I'lli:   rll<<  I  I  M'lDV    OK    Till     lll.iHili 


Ilif  I'liiTciils  mI  ii|i  1>>    till-  I'linliicl  ol'  iiii'lal   with  tlii'  Miliiit nstituci 

111'  till'  liilisrlf  jilircs. 

ir  «!•  ,-i.iini'ii  :i  LMhiiiiiiMii'trr  li,\  nii'iiii"  ni'  iiiiM|Hi|iM'i/.iilili'  i'l<'i-tii!.l 
with  twii  |i;irts  of  m  iIciioin  iiti'.l  iiiiisric  (tin-  <'iiinii/<<l  siii'toiiiiH  of  tl 
fiiiu".  it  "ill  III'  t'liiiiKl  tliiit  II  i-iini'iit  is  set  ii|i  >\  lifiicvof  n  wiivc  i>f  n. 
tiMctioii   |iiiNM's  liver  tlif  iniisclc   Iroiii  (Hii-  rnii   to  tlio  (itliiT.     Tin-   ]m 

wliirli  first  I'oritriifis  I mius  clfctrii'iillv   nrfiitivc  to  llif  rest  nl'  tlir  imisr 

lillt    ns    the    Mliv    of   i-olltllliMioll    |iiisscs    llloiiu'    it,    flic    "  ll<'!.'lllivif  V "    il 
rliascs   ill    tl «>\    (It    "llicli    tlic    \MI\c   still  Ir.l    >■       il.    «ll('ll    lIlP    >MIV('    li 

rciiflH'.l  till'   iiiiililli-  ol'  till'  strip,   iii'itlur  I'lid         llic   iniisrii'  sliows  ir 
ilitTficiH-i'   ill   |ioti'iiliiil.  so  flint    lln'  stiiiii;  I'onirs  Imck   to  ii    |iositioM 

ri'sl.     llowt'Xcr,  !is  tl iifrin'tioii   w.im'  icaclirs  flic  fiirf licr  end  of  |i 

llllisclc,  tllis  lellil  in   tlllll  lircollli's  nrifiltive,  illul   the  sflillj.'  s\\  intlH  ill   t 


^"lllllllllllllllllit 


a. 


A 


iMir.    .^1.      I>ia«raiii    I"    illii--Iiitlr    ih*     .Itvtit'ir 


»|.'i'.i'l    Ml     Ihi     «.ii.     if    inxjimn 


-hii'  "f   miiMlc    iiurari/Ml   s.ii  luriiii)    «!ini   'liiiiiilatrcl   iii    thr  tiu\    (/'I.      The   slia.lnl   imrliMi- 
Ihr  i.i.»ili..ii  u(  Ihi-   n.->!,itnilv.      Tin-  i.f.rliim  n(   tin-  nirvc  ■lia»n   l>v   ihc  .iifli  cli"n«  i.f   ihi-   Balv  . 
till    .11    i.ich    -!.i|ji     .III    -li"«u    .11    i1k>    riuhl    C.I.    '',    ■■.    aii'l    ■'••      i.Miir    l.i«i-.i 

npiiositc  (lirci'fioii   (  Ki^r,  .'i4>.     From  this  i-oiiip,iriiti\i'l\   >iiiipli'  cxperiiii. 

it  fiin  lie  si'i'ii  that  a  iiinsiMilar  i-ontrai'tion  wave  arises  at  the  eleefrotle  wli  u 

is  nejrative  first,  anil  that  the  iiioveiiient  of  the  striiijr  of  the  jralvaiioniefei  > 

most  marUeit     that  is.  the  iletleetion  is  frrealesf      wli-ii  the  two  eleetri"'  s 

are  applieil  at  the  extreme  enils  of  the  iiiusele.     When  tliey  are  liroii.  i 

eloser  to>;ethel-,  the  initial  ilelleetinii  hei es  lllllell  less  luaikeil  ;  ill  ol'  '' 

words,   the  aiiiplitnde   of  the   iiejralive   wave    is   i.'reate.st   when   the   t.  " 

interval    lietweeii    the    reeeipt    of   llie   eM-itatioii    at    the    two   eoiitaets  - 
'.'reatesf. 

The  applieatioii  of  these  laels  to  the  stiidx   of  the  initiation  of  the  I  • 

in    the    anrieie    reipiires    tliaf    we    shoiilil    eonsider    a'lotlier    proposil  ■ 

liMiiielv,   if  a    pail'  of  eonlacts  are   arraiij:ed   in   the  eeiiter  of  a   eiici  i' 

sheet    of   miisi'le   and    the   edye   of   this   sheet    is   stiiiiulated    at   diftV:  .\ 


nil     I'll  VMiil  iH,\    III      Till     Ml   MM'III   \  I' 


1M!I 


I       Is.  ilii>  iinipliliiili-  lit  ilt'ilri'iiiiji  III'  11  uiiKiiiiiiiiii-ii'i  I'liiiiii'i'li'il  with  till' 

|.    r  111'  fiPiiliirfH  will   lie  iimsl    |ii'iiiiiii!iir<'i|   \\  lu'ii    iIh'm-  iiif   niiliiil   to  tin- 

{I  iiit^  III'  Ntiiiiulatiiiii.  I'or  i  inlcr  tlicM-  i-oiiilitiiiiiH  it   is  i'\iili'iit   tlint  tlii' 

v'ti  ,iti'>*f  |)tiNHilil('  (lilTi'ifiirc  \\ill  i\isl   liilwi'cii  till'  iiiti'i'Mils  ii'i|iiii'i'ii  I'lir 

'I    ■    VUlXt'    fi<    ri'iirll    riM'll    cnlitiirt. 

I'r.ll'illlf  tlll'<t('  |ll'iliri|ili'>  in  niillil.  we  IMilV  imw  pliii'Ci'il  III  I'Xiilililli'  tilt' 
I : 'ill  HIT  iioiiiliiifi  III  llii  iiiiiiui  III  Ihi  III  III  Hii  III  III  Hii  siiiiiiiiii'iiiiliir  mull  : 
'  I      Wlii'ii    twii   I'lct'ti'tiili's    lire    ii|)|Mi<'<l    ill    ilitVi-i'fiit    |i  lints    lit'    the    an- 

11.  l'-,     lilt'     lllll|llilll<ll-     111'     lllll\»'IIH'l|t      III'     lilt'     stiilllf     lif     till'     U!ll\ill|ll|||l>tl>l 


.1    I. 


I'll     ill-ill'lllt'Ilt 


itcst     Will'll     til 


tllf    fit' 


IH'I'll    II.X'    t'lH'll     Ill-ill'lllt'Ilt     IS    iilClltl'st     Will'll     till'     lllir    Jllllllllt;    till'    I'll'l* 

ti.nlis  I'liiix  i'i)fi's   nil    tllf   siniiiiiiririiliii'    iniili'      Tn    iiiiiki'    this   rlfiir    tlic 

III  '  I  IMi'llI  III'  till'  stlini/  lilllst  III'  |i|intnui'i||)|ii'i|  ill  llii'  iiiiinili'r  alinvr 
.!•  -.  I  ilii'il.  till'  ii'siiltlnif  triiciiii;  lii'inif  rii'lril  jiii  t'li'i'trnriiiiliiitrriiiii.  I'iniii 
•I  -  r\|)i'i  iiiii'iits  w  itii  the  I'iri'iil.'ir  slit  rt  nl'  iiiiisi-lt'  tillinli'tl  In  it  is  t'viili'iit 
il,i'   till'  stiiiiiiliis  til  piiiilni'i'  tliis  itsiilt   iiiiist   liint'  misi'ii   in  tin-  iit'i<;li- 

Im.i  Im  ml  III'  till'  limit'.  \'2'  Ir  mil'  I'li'rtl'iiili'  is  |llilfl'il  nil  tin-  siliiiaill'ii'llllll' 
!  i.'l.    .iIhI   till'   ntlirr  I'li'i'tfnilr   is   lllnvi'il   alinlit    frnin    pljii'i'   In    plili'l'   nil    tin- 

mI I"',    till'    iji'lli  rlinll    lifillir    linti'il    ill     fili'li    lirW     pnsitinll,    tin-    I'lt'ft  I'liil" 

HI,  ili.'  limit'  will  !l!\\ii\s  III'  fiillliil  til  If  lii'UiitiM-  In  tllf  nthfl'  flfftlntji':" 
"hull,  lliiw  I'Xff,  will  lint  111-  till'  I'llsi'  ir  liiitll  I'lt'i'tliiili's  ill!'  Il|ii\i'il  illinlll 
M!i   ..IJii'i'   p.aits   nt'   tllf   Illllii'lf. 

As   Wf   sllilll   sff   ililllli'iliilti'jv  ,    till'   I'liri'flit    lit'   ili'linll    111'   till'   liflltiliH 

|i,''  may  In-  i-ffimifil  li\  I'lniiiii'tiii!.' a  uiilvaiiniiii'tf r  witli  vaiiniis  iiails 
C.I  'lir  liiii|\  ;  fur  i'\iiiii)ili'  with  tllf  I'itjiil  t'nif  liiiili  aiii]  tlif  It'fl  liiinl 
I.I  I.  Mil  till-  fli'vi  iiifiii'linurani  lluis  nlitaiiifil  aif  si'M'iiil  xxiivfs.  mif 
liii'll.  filllftl  tllf  I'-WiiM',  ran  I'lisily  lif  sllnwii  In  fnii'fs|)iilii-l  tn  tllf 
'  '  iii'tinii  111'  tllf  aiirii'lf  i  sff  rij;.  'Jtili.  1 1'  miw  wi-  fiinipiiit'  siirli  flif- 
'  '  iilinirranis  with  tlinsf  nlitaiiifil  wlijlf  I'liiitraftimis  nl"  tlif  aiii'ii'lf 
.;  •  |iinilui'fi|  liy  applyiiiir  iirtitiriiil  stiimiliitinii  to  varimis  jiiirts  of  it. 
ii  111  111"  t'niiiiil  tliiit  till'  iiitilifial  siiiiiilatfs  tlif  imnuiil  riir\t>  mily  wIumi 
'i  -liiiiiiiiitfil  part  is  ill  tllf  iifiu'lilinilinnil  nt'  tlif  siimaurif iilar  iinilf. 
I:  Mii'i'  wni'ils,  it  is  niily  w  lif  n  tlif  stiliiuliis  is  applifil  tn  tin-  siiiniliiiif- 
'i'  '  "iilf  tliiit  ii  fliiii'iiftfi'istif  I'-wiivf  is  nlitaiiifil.  Winn  tlif  iipiifiiilix 
!'■  siipi'iini'  viiji  fiixii  is  stiiiiiilatfil.  tllf  I'-wii\f  is  ilistnitfil  altlinui>li 
'lii'i'  wavfs  nl'  tllf  f Iff t tllf iii'tlini,'i'aiii   iiiiiy   lit'  imiiiiiil. 

Hy  taUiiii;  siiiiiiltiiiifinis  flcftriifiirtlinfriiiiiis  fi'mii  tlirfft  Ifinls 
■I  nil  tllf  aiii'iflf  iiiiii  fiiiiipariii^r  the  I'ffni'tl  with  that  nt'  ji  staiwliiiil 
'•lul  takfii  siiimltiiiifiiiisly ,  wi-  shall  liiul  liy  fxuft  iiifiisui-fiufiit  tli;il 
iiif  nf  niisft  nf  tllf  fxfitalinii  wiivf  nt'  tllf  luiriflf.  as  ir.>;isui'fil  in 
"iisliip  tn  tllf  !'  wiivf  nil  tllf  staiiilni'il  f!i'i-iri>f;iri|in>ri';nii,  is       i    ti'st 


!l 


til. 


uiitict  tit.n-i    tirtv.cen    thr 


itii.tUs    mitl    Kalv.)iHi!iii-Ui     .in-    .ilw.i> 


i"venient  of  the  nhailnw  ol    ihr  sIriiiK  almvr  llu-   li 
"Irctrk  tugaltvity. 


til-    tit    r.|UaI    pnti-lltial    .il    llu      Xv, 


I!t0 


Tin;  ciuci  i.ATKiN  OK  Tin:  iii.ihii> 


when  one  t 


IciMiimIc  is  (p\cr  till'  iippci'  (Mid  of  tlic  siiinaiiriciilai'  midc, 


thiit  ill  otiicr  i'(i:iiins  ot'  tlic  aiuifli'  il  alwavs  aiipcars  at  a  later  iiilciv. 
l-'mtliir  (IctiiiU  "II  lliis  siiiijcct  will  he  t'oiiiid  in  the  papcis  liy  Hystcr  .11 


.Mci'k"  and  in  Lewis.  iiiiiiinj:raplis. 


I' 


rf<|in'iilly.    Ml 


taking;   i'li'ilnicarili";;iiiiiis    freiii   ililTiiciit    parts   cf   tin 


lie.    It 


riiiiiiil  lli:il  iMTtaiii  cif  llic  iiirMs  s!iin\  siiiiill 


>  iif  |Hisili\  ity  1.1  lipw  tlic  liiii'  i>( 


pti'Mtlal 


'.liliu    tl 


II'    main    \\a\'i 


tKr.l    ull.ll    l,.itll    tl Ici-llMill 


if    m';;ati\ity.      'I'licsc    initial    ilctlritiens   air 
'i|    t'lniii   till-  siiiuaurii'iilar   iin<lt'      t'' 


s  arc  t'ai'  ri-innN 


iiii|.li',  hIcmi  tlic.  aic  I'laccil  im  the  auricular  a|  pcmlix  ;  liiit  tlicy  are  iicmi-  iiii'scnt  u 


._!.ay^-_,i_:If;: 


-^-^™  'y^_     _^:"r~, 

=re 

— «= 

T.^=^ 

;ff-^  [j::^  "^  -"■p'Tr" 

fT.'r:  - 

frrZ 

:      .'""-" 

- 

:*fcf-  jHf" .----■ 

.  f-^ 

■"■-- 

i 

7 

I'i«.     ^^        Sitim'l.iiit  ..ti^    tl(  iti.i..ii.li'.^;ianis  to     >ln.w      Ilu-     c;iii^r     f..r     cvlrin.ic     .Ml.  >  1 1.  it*.        1 

ill.l.cr    cm.-    .ni'     Ilhii     ;Iii     ;i|.|.<ii.Ii\     ;in.|    ill.-  l..»ir    ..m"     fLiiii    led     11.       Tin-    .lilt  I     ..r    mm 

iMliili.Mi  .  V  .■  1  I-  Mill  1.1  .li»a|.|..;ir  ill  III.-  I  ii;lll-l.;iii.l  .ii.)..  ll.li\  1 1.  i  li .«  ,iiih..i,T.ini.  l.i.aii-i 
I..1-.  ..i  111.  ..i.M -i.liv  lia<  I... 11  ,i;.-lii.|.  Til.  iMiiii-i.  .Iitl..li..ii  I /;  1  I  r.iM.iiii-.  .1-  'I.'  Ih' 
lri..il.,i    .litl..li..ii~    1  I   '    I      I.       1  l'|..lli    II  «i-.i 

I. lie  i.r  ihr  cli.it'...lcs   i..   |. laic. I  nil   tlic  siiiiiaiirii-nlar   iin.lc   ilsill'.      In   i.tlici    hmhIs.  iiii' 
taken   t'lmii   leail>*  at   a  ilisl.ainc   I'lmii   tlic  >iiiiiaiiriciilai    timlc  an-   in -ic  m    Ic-s  ci.iii|.. 
in  fiiMii,  l.ciii;.'  nia.lc  ii|i  paitlv  uf  llic  inaiii  ilcllcilii.ii  iliic  In  tin.  :iiii\al  iil'  \W  <'Xi'it;i' 
ami    |.artl\    uf   tlic   sccmlaiv    ilctlcctiiiii.i   ilcpcnilcnt    iipnn   extrinsic    iiitliiciices   acliiii; 

tl Iiitru.lcs;   that   is.   llic  clcclMiilc   picks  lip  I'lcclric   iliscliiir;;cs   fimii   .listant    ana 

iiiiisrlc  «liilc  tlicsc  arc  ill  a  cini.liti. f  cuiitractien   (  l-'i;^.  .'i.'i).     I'rinn  tliesc  c..n>i.l 

timis    il     t'lilliiws    th.-it    tlic    inicr\als    lutuccii    tlic    intrinsic    ami    extrinsic    ilctlcil 
sliuiil.l    lie    li.ii;;cst    in    ic.-iils   tliat    arc    t'artlicst    trniii    the    i.ii.lc.   ami    ;;r;ii|ii,.|ll>-    l.i  .  ■ 
less   as   line    lit'   the   ciiiita.ts    appricnhcs   the    mule,    until    hmt    Iliis   strnctiirc    tli. 
trinsic.  iletlei  tiiiii   is  nil   ImiL'cr   rcci.i.liii.     Sicli   lia~   I n    t'lniinl   tu   lie   tlic  case. 


I      ? 


'IIAI'TKU  XXII 

TIIK  I'ln     |i  (.OCY  .'I'    ;ilK  lIKAK'TIiKAT  (CciitM) 

THE  ORIGIN  AND  PROl'ACATION  OF  THE  BEAT  (Cont'd)— 

FIBRILLATION 

Mode  of  Propagation  in  the  Auricles 

I'ldin  tlic  mass  i>(  ex 'nlciicc  we  have  littli-  ilmilit  tliat  tlic  licaitlioat 
11  ijiiiatcs  ill  tlic  siiii'Miiiiciilar  iimlc.  anil  tlic  (|iiislinii  now  iiii'si'iils  itsi-lf 
,i--  tu  liou  till'  lii'at  is  jiriipayati'il  hmt  tin'  ri'maiiiilcr  of  tlir  aiiiicli's  and 
I  lo  till'  \i'iitrii-lcs.  Hcu-anlinsr  tin'  piii|iayatiiiii  of  tlii'  licat  nvi'i'  tlir 
iiiiiii'li's.  two  ivissiliilitii's  cNist:  I  I  it  may  sprcail  iinil'nrmly  over  tin' 
iiMi-M'iilar  tissiii'  of  the  aiii-inilar  wall  until  it  rcarlics  tlu'  aiirii'iilo\i'nti'i('- 
iii.ii-  iioili'.  or  i2'  tlit'i-c  may  l.c  laiil  ilmvii  lictwci'ii  the  siiioaurirular  and 
III'  amiciilDvcntiii'ular  node  a  spci-ial  strand  of  lii^^ldy  condiii'tiiiir  tissno. 
I'  is  no  ai'!.'umi'iit  ajrainst  tliis  si'i'ond  possiliility  that  wi'  slimild  so  far 
li:i\r  liccn  iiiialih'  liy  liistolo<;ii-  nii'tliiuls  to  dilTt'i-intiatc  any  sui'li  stiui'- 

Thill'  is  considcralili"  praitii-al  importani-c  attarln-d  to  the  solution  of 
tin-,.'  i|Ui'stions.  jiai-tii-ulaily  witli  n",'ai-d  to  tin-  rausi'  of  ci'itain  fyjii's 
■■I'  r.iriliai-  ai-l-liytlimia.  siicli.  for  example,  as  that  known  as  iimldl  rhiitlan. 
T!  i|N,  it  is  evident  that  if  the  heat  is  transmitted  uniformly  over  the 
I  ';~'-nlar  tis:,iie  of  the  auriele.  then  the  whole  aiiriele  will  have  eon- 
!  irlcd  hefor,.  the  heat  has  reaehed  the  aiiiieulox  ent  lieiilar  liundh'.  Iiy 
■  !  I'll  it  is  then  transmitted  to  the  \  eniiieles.  On  th"  other  hand,  if  the 
■i.ii  should  ti'axel  hetvveen  the  twn  nodes  liy  speeial  eondiletinjr  tissue, 
t!ii>  Ihe  im)iulse  will  ha\e  arrived  at  the  aiii'ieiilov  entrieiilar  nnile  lie- 
i  '!■•  the  auriele  has  eonlraeteil.  As  a  matter  of  faet,  it  is  not  quite  settled 
>'l  a-<  to  wliieli  of  these  two  views  is  Ihe  eori'eet  one.  althoil^dl  the  halaiiee 
'■>  ■vidence  serins  to  favor  the  former  that  is.  that  the  wave  is  transmitted 
•  '!"rmly  over  the  iiiuseular  tissue  of  the  auricle.  (Lewis.~i 
The  iiiethoils  einployed  in  altaekiiii:  the  prolilem  have  heeii  esseiitiall.\- 
"  '■  same  as  those  descrilied  ahove,  One  of  them  may  lie  called  the  direct, 
I  '  cither  the  indirect.  In  the  former,  a  series  of  pairs  of  contacts  is 
I    ■•■1(1  on  the  auricle,  ejieli  pair  Iieiiiu-  in  a  radial  diieciion  to  the  sjuo. 

iry  process  arrives  at  that 
il 


I  ular  node.    The  time  at  which  the  excitat^ 
lift  of  each  pair  \\  liieh  is  proximal  to  ih 


le  sinoiiiirieul.'ir  noile  is  accil 


lUl 


Ill- 


;  III    ciifri  i.v'i'iiiN  Ml'   rill    I'.i.ihiu 


liili'ly  ili'lfiiiiiiiicl   Ir  nil  Ihc  Lriilviiiiiimcliic  iimihiI.      Tlir  cxai't   ilistaiUM'  Ih 
twiiii  the  ((iiitai'l  aiiil  till'  siiioaiirii'ular  iumIc  is  then  iiicasiircd  and  (vmi 
llic  ilata   till-  a\i'iaui'   Iraiisinissioii   tiiiii'   is  cstii'ialcil.      I'roiii   Ills  vcsiill 
l.cuis-  (•(iiii-ludc's  tliat  tlic  traiisniissidii  rates  aii-  iinil'oiiii   rroiii  tlic  iiml' 
t(i  all  parts  ut'  tin'  aiiiii-li'.  with  llic  i'\fi'|iiiciii  ol'  tlic  sii|)('i-i(ir  \t'iia  s .  \  ,i 
ill  wliicli  llic  late  is  (•(iiisidi'ialilv    lower.     One  tliciiisaiid   iiiilliineters  )" 
second   re|iiesehts   \i'iv    t'aiily   the  aveiaire   rale  at    wliicli   llic  cxcitatiHi 
wave  traxels.     On  tlic  utlicr  liaiiil.  K.\stcr  and  .MecU'  state  tlial   the  \\a\' 
is   |(rii|iai:ated    tliinU'rlKiiit    the   sinus   ikmIc.   and   tiiat    it    sjireads   to   IIm 
ciiiiti<riiiiiis    \eii;e   cava-   and    to    the    auriculnv cut licular    inxle    with    ecu 
sidcralile  rapidity,  reaeliintr  the  nnnith  of  tin-  superior  vena  cava  in  ''.I'l 
second,  whereas  its  passa^'c  to  Ihe  auricle  itscli"  takes  O.O'J  second.     Tliere 
tlicicl'orc  a  delay   in  the  passajje  nl'  the  wave  to  the  auricle,  which   iinl 
catcs  that   the  excitation   iiiiist  spread   to  the  aiiriculoveiitrieiilar  node  1" 
tnrc  iii\ol\iiitr  the  liiilit  allium.     'I'liese  authors  conclude  that  "this  lea'l 

til  the  iiie\italili iicliisinii  lliat  the  cardiac  iiiipnlse  spreads  to  the  vei 

tiicle  and  to  the  rii;li)    auricle  hy   ilitTerent   paths,  and  does  mil    pass  l 
the   \eiitrie|e   ihiipiiuli    the   auricle,   as   ordinarily   stated." 

ill  the  s lid.  or  indirect,  nielhod,  the  onset  of  the  negative  WM\e  I'loii! 

dilTcrciit   leads  in  the  auricle  is  compared  at::aiiist  a  standard.     l''or  tl  ' 
standiird   Kyster  and   .Meek  h:i\e  used  the  record  of  the  mechanical  s\ 
lole  of  the  .iiiricle.  Iiiit   the  iiilcipretat  ion  nf  the  result    is  extremely  lii 

licillt  on  a Hint  of  the  rat'    at  which  the  diaiifres  are  oceiirriliir.     I,ew  i 

nil  the  other  hand,  has  used  the  standard  elect rocardioLrram  for  pirp.isi 
of  cniiiparisnii. 

Mode  of  Propag~ation  ot  the  Beat  to  the  Ventricles 

After  iiMchiii^.'  Ihe  aiiriciiloventiiciilar  node,  the  heat  is  t ransmitteil 
the  vcntrii'les  aloiij.'  the  aiiriculoveiitrieiilar  limidic  a  fact  which  has  l'> 
most  clearly  deinoiistrated  liy  the  experiments  on  In (irl-hlix'h-.  We  have 
rcidy  Mcii  i  fiajre  174i  that  althoiiirh  each  ehaiiilicr  of  the  licarl  ni 
turtle  or  froii  has  a  rli\thm  of  its  o\\  p.  this  is  mnidi  more  pronounced 
the  \eiioiis  end  of  the  heart,  and  when  the  transmission  of  the  lieat  to  ' 
\eiitricles  from  the  auricles  is  olistiucted  or  hlockcd.  as  liy  compress' 
or  partial  i-nttin-i  at  the  aui'iciilo\'enti  icillar  .junction,  the  xcntricl- 
after  coiiiiiiir  to  a  standstill  fiu'  a  time,  siihse(|Ueiill,\'  contract  with 
i'h\thiii  wliiidi  is  ciitii'el.\    iiidependcnt   of  that   of  the  auricles. 

In  the  mamiiialiaii  heail  the  same  results  may  lie  olitained  h,\  ari'a> 
iiii.'  a  clamp  so  that  it  cumpresscs  practieall,\  nothiiiir  hut  the  auric!; 
\entricnlar  liundle  i  Krlaiiucr.  i  If  the  compression  is  extreme.  ' 
rhythm  of  the  ventricles  is  (|iiite  iiidependcnt  of  that  of  the  auricles,  1 
if  it   is  onl.\    partial,  the  \  cut  i  icular  systoles  follow    rcirularly  e\ei',\    - 


Tiir.  I'livsKii.odv  oy  nil:  iii:Aicriii:\'i 


mn 


=-3 


111,  tliiid.  X     loiirth  iiuiiciiliii-  ( triictioH.     If  iil'tiT  siicli  ii  (•i)in])l('1<>  of 

iitiiil   liPiirl  lilock   liiis  hci'ii    instituted,   tlic   cljinip   is   iciiinv.'d,    it    will 
-iially  lie  foiiiitl  tluit  tlic  licjirt-lildck  disMppcais  .-Mid  tiic  iiiiiii'iiliii'  and 


itriculi 


ir  contriictioiis   t'al 


I'k    into   tlicir  usual   scciui'ii'mv 


Th^ 


.'■laiico  (if  this  (lisc(iv('r\,  apart    fi 


loni   its  |)liysi<il<i<;i<-   niti'icst,  rests  in 


]■.«' 


fact  that  it  is  irncihi  iliiplicitli  (J  in  cliiilriil  r.ijK  ri(  iici .  If  the  ])ulse 
lUfX  (d'  the  radial  arterv  is  eoinpared  with  that  of  the  .;ujrular  vein 
el-taiu  types  of  liearf  disease,  it  will  lie  found  thai  tie  auricle  is  lieat- 

n   inanv  of 


two  or  three  times  more  i|iiii-kl\-   than   the   \  eiitrieles.      I 


ihi'se  cases  it  has  lieeii  found  on  auto|)sy  that  definite  lesions  id'len  syphi- 


le  ill   nature  inxtdve  the  aiiiieuicx  ('Utricular   Iniiidle.      In   otl 


ler   cases. 


li.wi'ver,  such  lesions  haxc  not  heeii  discovered.  Sometimes  the  tnindie 
i-  Ml  severely  diseased  that  the  liloidi  is  complete,  the  ventri(des  coii- 
t'actiii!,'  (luile  indepelidenlly  of  the  auricle  (Stokes  Adams  syndrome.) 
|i:  such  cases  it  is  assumed  that  the  lieal  oiiu'iiiates  in  the  unin.jured  jiart 
..I  llic  liundle  below  the  seat  of  the  liloek.  It  should  lie  pointed  out  here, 
l.nwcver,  that  all  eases  oi'  slow  juilse  in  the  arteries  are  not  necessarily 
il>  ii.iidcnf  upon  heart-lilock,  Init  ma.\  depend  upon  a  slow  heal  of  the 
ii!!i!i!c  itself.     This   is  called   hraihivarilin. 

SiiiMctinies  after  c(im])lete  destrucli  iii  of  the  aiiiiciiloventricul.ir  huii- 
lil.-  ilie  heat  continues  to  lie  transmitlcd  lo  the  \i'ntri(de,  and  coir  -scU- 
til--  transmission  has  sometimes  lieen  oliserved  to  lie  upset  li\-  lesii  lot 
^liV- 'liiij;  the  Iiundle.  The  explanation  of  Imth  of  these  e>;ce|)tiop;il  I'c- 
vir's  almost  ceitainl.N-  is  that  the  riu:ht  lateral  connection  descrilicd  aliove 

|i.ii:r  I.S4)  is  .scrvinyr  as  the  main  pathway  of  transmission  for  the  heat. 
I  111'  facility  of  conduction  throutrh  the  auriculoventriculac  Imndle  is 
>'i'  jri't  to  alti'iaticin  liy  the  imjuilses  passiiij;  to  it  aloiiLr  the  \ay.ls  nerve, 
p.i'iiiilarl.v  the  hd't  va«>us.  It  can  also  lie  altered  hy  ccitaiii  druses, 
'-  ■ 'iMlly  dijiitalis  and  strophanlhiii.  The  dear  demoiistiiition  that  it  is 
■■  ■  ■•  this  liundlc  that  the  licat  is  transinitlcd  is  st-ronu;  evidence  in  favor 
■!ic  iiiyofjeiiic  hypothesis  ( pajrc  171  i  coneeiniiii;  the  I  ransmissioii  of 
!i      iirartheat,  hut  it  does  not   necessarily  disprove  the  neiiro^renic  liypoth- 

loselv 


iiir  histotoiiic    invest iyat ion    has   shown    that    ih 
iiliided  liy  iiii  intimate  jilexus  of  iier\i'  lilieis. 


le    liundle    IS   ( 


Spread  of  the  Beat  in  the  Ventricle 

">nir  the  impulse  has  lieeii  transmitted  liy  the  liundle  into  the  vcii- 
''  '^.  it  spreads  aloiiy;  the  many  liraiiclics  into  which,  as  \\v  have  seen. 
''  '  '  I  main  di\isions  of  this  liundle  separate.  The  tirst  i)art  of  the 
'  • '!'iiciil;ir  musculature  to  conli'acf  is  ttierefore  'ucated  near  the  trr- 
I'l'  'inn  of  these  liraiiches.  at  the  i>apillary  inuscles.  That  these  should 
'•"   'i.ict  hefore  the  rest  of  the  muscle  of  the  \ ciitricles,  lias  an  oli\ioiis 


f 


jj 


l!t4 


TIIK   (IHt  TI.ATIKN    (IK    Till:    I!I,Oil|> 


si<rniliiiiiii'i"  in  ('(imH'ctiiPii  wiili  tlicir  t'liiict ton  t>(  tin  iiiir  tlic  clionlii 
londiiit'ii'  Ml  iis  t(i  ])ro\i'nt  :iiiy  Iniliiiiiir  nf  tin-  lliips  di'  tln'  aiii'iciilovon 
Iricnlar  v:ilvc  intu  tiic  jiiiiii'lcs  w  lien,  at  tlic  lu'iriiiiiinu'  ol'  llic  iircsplix  iriiii. 
pcriml,  tlio  liiirli  inliavcntiiciil.ir  incssin'c  is  liriiniilit  to  licar  on  llirr 
iliulci'  siiri'ai'i's.  Al'tcf  stattin^  at  tliis  |iiiiiit  in  the  vtiiti'icio,  tlio  ••mi 
tiactiim  \va\c  si'diis  to  sjH'cail  lartlicr  tlinniirli  Ilic  vcniiicular  iniiscio  a! 
a  fairly  unii'orin  rate. 

Invcstii:a1i<iii  ul'  tliis  ]ir(ililcin  \<y  means  uf  llio  •lalvaiioniclcr  lias  Iiimii 
ti'rlinicallv  a  \('i\  ilillicrlt  iiiattrr.  and  tlit"  details  (if  tlio  fcs('ai(dics  li.\ 
Lewis  and  liis  piipils  lia\('  not  as  yet  lieen  |illlilislied  in  full.  Aecdrdinj 
111  till'  iirerniiinar\    ennininnicat  imis  at   hand,  Imw ever.  '   it   apjiears  tliat. 


■0Jl6/ 


I'l;;.  '(,.  |Iml;i,,im  -i'  <  \im  t  I'll,  tit  l.\-  I,<ui^  -li.uiiiy  till  tii:n»  at  ulii.Il  the  r\titattnn  w 
.•ipl"  If. I  .ill  till-  It.. tit  .1  111.-  li.,irt  lilnni'  t'i  III.  iii-lr..lM.  ii  A'  in  I..1.I  II.  A'., I,,  liKlil  :i|.i  • 
ill-.:    /'./'./...   .1.^.  iiiilini;    l.iai..li    .■!    I.li    .■..i..ii.ir\    ,111. ty.      ll-'iii-ii    Tlti.iii.iv    l.iwisl 


If    :' 


uhen  niin]iiilari/.alile  elect rndes  are  |ilaee(l  at  vafimis  parts  "f  tlie  mit' 
aspe.'t  (if  the  veiiti'iele,  and  einiipai  iscni  made  uf  the  iiiomeiits  at  whii 
the  cai'iliac  impulse  aiiixi's.  as  jiidycd  liy  the  a])peai'ancc  uf  the  exeiK 
timi  w.-ivi'  relative  tn  li  in  a  standard  electnieardidiiram.  it  has  Ih' 
fiiiind  that  the  time  (if  arrival  hears  mi  relatimiship  to  tlio  aiiatoniic  a' 
ranirement  of  the  muscle  Inindles  of  the  ventricle.  It  arrives  early  ai 
siimiltaiKHiiisly  over  an  area  of  the  surface  near  the  aiiteiior  attaelmii  i 
of  tlie  \\all  (if  the  ritilil  ventricle.  It  arrives  Into  at.  the  Iiaso  of  tlio  ri'-'i 
ventricle  and  in  the  part  near  the  poslciiur  inl  rav  cut  ricular  ^roo\ 
Ilistdloiric  examiinitioii  has  shown  that  the  Iiranches  of  the  lifrlit  divisi. 
of  the  ar.rieiiloveiitrieiilar  Imndle  are  iiiost  (losolv  connected  with   li 


k 


Tin:  I'livsKii.ocv  (IP  Tin;  im:.\uti!i:at 


195 


phifi-  wluMc  tlic  wiivc  ■■urivf-i  ciilifst.  Sciiu'wlint  (litVciciit  ri'siilts  arc 
..I'liiiiicil  iVoiii  till'  lift  M'litrirlc.  Iiiit  iiu::iiii  tlicy  Jilc  ilipi'liili'iit  lliinii  tlif 
IrhltilPllsllil)  (if  tlic   pMit   to  tlic   I'lii-Uiiijc  tiliCIS   I  l'"iir.  .'id). 

FIBRILLATION  OF  THE  HEART 


-'I 
■1 


Ventricles 

Tlic  even  si)i('a(l  nf  the  wave  of  coiit lai-timi  hmt  tlic  licait  ijcpcnds  <mi 
tlic  uiiil'diiii  cNcitaiiility  ot'  tlic  iiiiiscnl.ir  fillers.  If  certain  nf  tlic  nniscii- 
l;ir  fillers,  or  liiiiiillcs  lit'  liliei-s,  lia\c  a  iriralcr  iir  less  cxcitaliility  tlian 
i.iImin,  tlicii,  nlicri  tlic  stiiiiiiliis  to  cmitract  arrives,  it  will  not  produce 
,1  unif'oi-iii  ciiiitraction  oi"  nciyliliorimr  liiiiiilles.  jniil  crorilinatcil  action  of 
ilie  eariliac  iiiiiscnlatiirc  w  ill  irivc  place  to  a  confuse, I  nioxenient  in  wliicli 
i'.hIi  i>art    of  tlic   lieart    is  cont laetiiii.'   inilepcni|eiitl\    of  tlic   rest.     'I'liis 

lil'i-illiiliiiii.  or  liiliriinii  rnrdls.  as  it   is  calleil,  can  lie  proilii I  liv  a  larec 

',:riety  of  c\pc|-inielltal  liictlioijs.  siieli.  for  cN.iniplc.  as  liy  st  iiiililat  iliu' 
llir  ventricles  with  indiiccil  clcetrii'  sliocl<s,  or'  liy  liu-atjon  of  a  lariic 
hi.iiieli  of  tile  coronary  artery,  or  li\    tlie  in.jcction  of  lycopoilium  spoics 

ii'u  the  coriinai\-  cifcnhilioii,  or  li\    nieclianicil  stiiiinlati f  ilie  lieart 

it!  ihe  rceion  of  till"  aiirienlovcntriciiliir  lininllc. 

I'ilirillation  of  tlic  ventricles  is  mitloiilitcilly  ;i  cminion  c,-iiisc  of  dcatli 
II!  man,  for  of  course  tlic  confused  nioveiiieiits  make  tlic  vciitiii-li's  iii- 
iMliiililc  of  coiitraetiii^'  on  the  contents  of  the  heart.  It  is  a  comlitioii 
V.  hii-h  can  pi'olialil.v  never  lie  recovered  from  in  the  liiirhcr  animals,  Imt 
1'   i-  iif  interest   that   the  ease  with   which   it    is  set   U|)  as  the  result   of  the 

■  M'l'iii-ation  of  an  electric  stimulus  varies  to  a  marked   dcL'rce  in  difTcr- 

■  '!t  Miiiiials,  and  that  in  those  hearts  in  which  tilirillation  cjin  he  clic- 
'■'i  "nl>  with  dil'ficiilt.v,  recovery  c;iii  usually  he  cfTeetcd  either  hy  slop- 
!'  -  the  heart  liy  means  of  cold  ami  then  allow  in-,''  It  to  heat  ajiain.  or 
'.'-  tlie  administration  of  cpiiieplirinc.  ( tf  the  lie;irts  in\  cstiL'alcd  in 
th^  way,  that  of  the  rat  li.'is  lieen  t'ouinl  to  he  most  resistant  to  stimula- 
'  '  i;  then  in  oi'der  come  those  of  the  raliliit,  the  c.it.  the  doi:.  ami  the 
I'iM'.     There  is  L'ood  reason  to  lielicvc  that   the  heart  of  man  is  rcadil.v 

■  '  '  -'ill.  l''ilirillation  of  the  ventricle  is  iindouliledly  the  main  cause  of 
'I'.'h  ill  most  cases  of  I  hftnii  tdiiiii.  Curiously  ciiounh,  however,  it  has 
'"'"  -'ated  that,  whereas,  a  current  of  ordinaiy  intensity  i  L':!00  volts 
iiii' ' :  atin;,' eiirrent  1  |irodui'cs  ventricular  filirillation  in  the  heart  of  ecr- 
'■ii     •■(  the  lower  animals,  at   least    in   that   of  the   horse,   a   verv   nnicli 


•1'  current  does  not  d> 


Hid  iiiav  indeed 


cause  V  I'll!  lieiilar  fitnil 


I'liM'i   jiroduccd   liy    a    more    modeiale    vollaec   tn   disiippcai'.      I'nfnrtii- 


:it.'Iv.  1 


loucvcr,  these  stronjicr  cuircnts  produce  irrcpar 


uilc  (faiiiMLre  III 


i-  ji 


1!m; 


llll     tIKil  l.X'lliiS    111'     I  mi:    lll.lini) 


*        'J 


llio  (•(■iltliil  iKMVi.iis  s.vslciii,  s.i  thill  the  liK'llliiil  of  jipplyiilJ.'  stnilif;(M'  cllr 
iciits.  I'vcii  wen-  it  iVtisilili-  to  do  so  (|iiickly  ciiou;.')!.  would  lie  ot'  im 
tlicriiju'iitic  xiiliic  in   rcinoviiis,'  (ilii'illalioii. 

Till'  disjililioilitiiiir  results  tliiit  Iimvc  followed  the  lejieMted  iltteiii|it- 
to  |-esils<-it;it«'  Jicrsoiis  Uilleil  ;ieeidellt.lll\  li\  cleetlie  slioeUs  is  lllldolllil 
eillv  dependent  iliion  the  fiiel  that  in  the  lieait  of  man  it  is  iiii))ossili!. 
to  Iniiiu  liaek  the  normal  heat  after  tin-  ventricles  liave  lieeii  thrown  iiit.i 
tihrillation.  I'ilirillation  of  the  veiitriele  is  also  the  cause  of  the  sudden 
cardiac-  failure  oi-curriii<_'  when  lilood  clots  or  eml)oli  cause  a  lilo(d\a'ji 
of  the  eoroiiar.v  circulation  lit  is  sometimes  the  c-ause  of  aiia:inii  ))er 
tolls,  for  example  ,  It  must  also  he  rememliered  in  eliiiieal  l>ractie. 
liiat  meehanii'al  stimiilalioii  of  the  vi'iitricles  ma\  i)iddiiee  lihrillalioii.  s.. 
that  in  attemiited  resuscitation  liy  cardiac  lllas^a^e  c-are  should  he  taken 
not  to  ai)i>lv  this  too  viirorouslv . 

Auricles 

Althoimli  \eii1ricular  lilirillation  is  seldom  recovered  from,  it  has  In 

clearly  shown  in  recent  years  that  lilirillation  of  the  auric-les  is  relative  I'. 
coiiimoii  and  that  it  is  hy  no  means  iminediately  fatal.  Indeed  it  is  on. 
ccf  the  most  commccn  of  the  ehrcniic  cardiac  disicrdcrs  in  man.  Auricuhr 
tjlirillatioii  can  lie  prodiic-cd  c>xperiiiientall\-  ii\'  the  ai>plieation  of  : 
stroiitr  electric  stimulus  to  the  auricles.  If,  however,  a  weaker  stimuli;- 
is  .-ipplied.  the  auricles  do  not  c^i,  into  t>  jiieal  lilirillation,  liut  come  '- 
licat  at  a  very  rapid  and  riL'ular  rate,  perhaps  three  or  four  liundred 
minute.  This  c-onclilicin  is  c-;illec|  ••auricular  llutter."  and  is  cpiite  f i  • 
c|iientl\    oliser\-eil    in   tlr,'  clinic. 

■/'//(    iiilhiitici    III'  (iiirii  iiliir  filirilliiliiDi  (iiid  Ihillir  mi  Ihi    hull  iif  llii    r. 
Iriih    is  iiii   c\treiiiel.\    iniiioilaiit    one   in    eoiiiicclioii    with    tho    irrc«^ul.:; 
ities  ccf  the  heart   iiliser\fd   in   man.  and   this  iiillueiice   in   most   cases     - 
explainecl   liy  i-oiisiderincj  i|      the  narrowness  of  the  jiatli    i  in  the  iiiiii 
uloveiitiicular  liiindlei  alccii'j  whic-h  the  impulses  ha\e  to  travel,  and     - 

tjic  \ \iiic_'  conditions  of  exc-i1aliilit>    of  the  \ cntricnlar  muscle,  depci, 

iiiir  upon  the  c'xislenc'c  of  the  refiac'tory   phase   i  pa'j:e   ISdl. 

In  auricular  Ihitter,  when  three  or  four  hundred   impulses  per  miinr 
;ire  ]i;issin'^  aloii'j   the  liuiidle  to  the  ventricle,  the  contraction  ])roduc  ■   ! 
li\    the  first   one  will  scarcely  have  started   '  "fore  the  second  and   inn; 
diatcly  succeedincj  ones  arrixc,  so  tlmt  the  ventricle  will   lieat   at   a    r, 
Unit    is  much   less  than   that   of  the  auricle,  unci  a   condition   simulaii    .' 
heart  hlcick    will    Icecomc    cstalilislied.      Tlie    cliarac-t:':istie    feature    wh 
distiniiuishcs   this    from    true   he,i!l-lilock,   however,    is  the   fact    that    '    ' 
ventricular   r.itc   is  iilmri    noriiial.   whereas   in    true   lioart-lilock   the    r 
is    miic^h    lielciw     normal.      H.\     means    of    the    eleetroeardio<iraiii    or 


Till:  I'liVMoi.ddV  UK  -iiii:  iii;.\iriiii:Ar 


1!)7 


-# 


i|\  >|iliys;Mii>)ir!ii)liic  trn<'iiii.'s,  it  I'liii  alsci  \n'  sIkpuii  tliiit  tlic  .•nii'icle  is 
.itiiiir  with  ixTlVrt  iru'iitiiiil.\  iiltli(pU!.'li  \«'i\  rfii)iillv. 
Ill  ciuriciiliir  (iltrilliitidii  tlic  vciitiiclcs  uliviniislv  will  rcspdiid  iit  ii  \r\-\ 
ii-mihir  rate  Id  iIii'  iinpiilscs  tiMiisiiiitti'il  to  tlit'iii.  iiiul  tli<-  •'luririilar 
iiiliiirtidiis,  if  cxaniiiiCil  liy  llic  iiu'IIhkIs  almxi'  (IcmmIIkmI,  will  slinw  in> 
yillar  .MMilU'lii'c.  I''iiit]it'i'  details  dl'  tlie  iiK'tliiid  iif  cliritiny;  t'lcsi"  ^iu'iis 
ill  lie  (li'scM'ilM'd  later  (i>aj.'e  J(i(>i. 


i   I 


■a 


fi       i 


CIIAI'TMU  XXlll 

TiiK  r.i.(Mii>ii.(i\v  IN  Tin;  autkk'Ik.s 

THE  PULSES 

lirtiii  ninu  In  llif  |i|i\  si.'al  law  s  that   unvili  lln'  "'ircMliiti I'  tlii'  IiI'mmI, 

\\v  iiia\  iMiw  (■■iiisiilci-  ilii-  |(  iiipnfaiy  c-liaii'j,i-<  |iniiliici'(l  in  llic  IiIimmIiIuw 
ill  tlir  aiti'iii's  l.y  iMi'li  s.vstiilii'  (liM-liaiirc  Tlic^c  clianucs  u'"  niidci'  tin 
LTiiiTjil  tciiii  111'  llir  iMii-.i's.  (if  wliii-li  llirci'  iiia\  Kc  ili^tiii'-'uislicil:  '1 
till'  |)ic'^^ini'  iiiii>c',  nr  till'  imKatilc  iiici-raM'  of  |iicssiiro  prixliiccil  li> 
cacli  hcarllirat  i  sec  pauc  12'-.  'J'  iIh'  vclM.'iiy  \m\>v,  ,<\-  puUatilc  accl 
ci-aliciii  III'  \clnrity:  ami  ^ :! '  tlic  |ial|(alilt'  |iii1m'.  nr  tlir  imlsatilc  cNpaii^i'Mi 

i.t'  till'  ualls  i.r  till'  M.i.i.I  \.->-.(U  |iru(liii-i  il  l.y  till'  MhMfll  i-liall'_'r  "f  M I 

|ins-,un'    ill    tlicir    iiilrrinr.      'I'lic    'jciH'ral    cliaractfiislics     cil'     tlic     tiin  .• 

pilUfv   ail'    till'   Naiih',    riM-tain    iVatiiri's   Li'lm;    Imw  cN c,'    l ■>'    |>n.ii(Hllii'c.| 

ill  oiii'  lliaii   ill  aiMitlii'i'. 

General  Characteristics 

Rate  of  Transmission  of  Pulse  Wave.  Tin'  rati-  n|'  iiansiniNsion  m 
tlic  iiiiNi'  \\a\r  call  I"'  i lit iiiiii iii'd  l>\  takilitr  siiiiilltaiii'Dlls  trai-iliu's  ol 
llir  |iiiKi's  I'liiiii  two  far  ili^taiit  j'art-^  <>(  tlir  arterial  systriii  iilmij;  >vilii 
ai'ciiratr  time  traciiiL's.  Fmiii  i-iMi.rds  ii-f.  Fi-.'.  !ls  i  taki'ii  frmii  tlic  ii|icN  '■'■ 
llic  raiiiliil  anil  ladial  artiiirs  \\  i'  call  ilctcniiilii'  Imw  Imiir  it  tflkcs  t'c" 
the  lic'^iimiii','  of  tlic  puUc  \va\c  t"  travel  In  tlie  radial  artery  i'riini  lli' 
liniiil  ill  the  amla  finui  which  the  earntid  artery  spriii-.'s.  We  shall  lii"! 
that  it  takes  ahniil  niie-teiith  >i\'  a  sceniid,  \\hieli,  cniisidel-in','  the  leli'^'!- 
nf  ihe  artery  iiiMihed.  wmild  wnrk  mit  as  a  Irjiiisniissinii  velncity  d' 
aliniit  se\eli  iiirteis  per  secnlid  nr  aliniM  seNcnteell  iniles  ail  liniir.  Tl  ■■ 
pulse   thenrnre   travels   alniis.'  the   Idimd   M'ssels   at    a    lllliell    '.M'eilter  spei  .| 

than  the  hi 1  it>ell'  is  iiiuvii  ■„',  this  lieiM>r,  as  we  shall  see  iiiinu'dialel ;,  , 

ahniit    II.-")    llielelv   per  srcniiii    ill    the   lander  lilniid    vessels 

The  pulse  is  a   wave  nl'  sudden  increase  in  jtressiirc  and  velncity  p.i- 
inu-  aliiiiu'  a   stream    v\liich    is  tlnwiiiir   in   the  same  dii'ectinii    with   a   <■•  ■ 

tain   iiinre   periiiancnt    pressure  and    velncity.      A   simple   phv  sical   e\p 

meiit  ma>    serve  In  make  this  dear:    1 1'  the  first  of  a  nivv  nf  hilli.'ird  hii    - 

he  tapped   v\itli   tl lie,   the  end   halls  will   lly   niV  while  the  nthers  .■:    • 

liinviiiLr  slnv\ly  alniiir  in   the  direetinii   nf  the  sirnke.      Kiich   hall   lieenii    > 
a leratid   li.v  the  hall   heliiiul  it,  and  imiiarls  lis  inlhieiice  to  the  I-  I 

I'JS 


■riii:  iii.oni)i-i.u\v  IN  Tin;  Men ;kii;: 


1H9 


I" 


rnml.     hi  iillici'  utiids,  ;i  piilsMtilc  iii Itiiiticpll  nf  \cliicity  is  piiidiiciMl 

.  ;i  pnlsfitilc  <'liilll!.'<'  ill  Iin's>iii'r  lirtxM'i'ii  ciii'll  two  l);ills.  Till'  cxisti'liri- 
'  II  pulse  «avc  L'oiiitr  in  tlic  siiiiic  (iirrcliini  liiit  (|iiicl<or  tlinii  ii  iimviiiu 
liiiiii,  ill"  tlniil  Clin  iils.i  lie  illiistnitcil  !iy  dlisiTvin;,'  tlic  \vii\ps  tiiivclins; 
iw  II  11  stii'.'iiii  wlu'ii  ji  st(iii(>  is  tlii'iiwii  intit  it. 

Till  Iciifilh  of  III  i>iihi  irnri  is  siicli  tliiit  tlic  Iicsriiiniiiir  of  it  lias  ar- 
,(i|  at  tlic  pdipliciy  (if  the  fiitciial  system  licfoie  llic  end  lias  ilisap- 
•.irrd  friiiii  tlie  lieuiniiiiii;  of  tlic  aorta.  This  is  iiiiixntaiit  to  rcineiii- 
Ki  I-.  fill'  it  is  a  coiiiiiioii  iiiistaUc  to  think  of  the  wave  as  hcinu'  a  local 
,,ii,'.  The  ilcleiiiiiiiation  of  the  Icnirth  of  the  pulse  wave  depemls  upon 
llie  follow  iiii:  ci|iiation:  I,  \'T.  \vli(rc  I,  cijiials  the  leiijrth  of  the  jiiilse 
\:i\e,  V  its  velocity  of  t  iaiisini>sioii.  and  T  its  duration  at  a  i,'iveti  i>oiiit 
ill  ilie  artei-y.  I'lichT  in-dinaiy  eiiciiiiiNtaiices  I,  would  usually  work  out 
iV.iiii  :i.2'i  to  4,r>  meters. 

The  rate  of  transmission  of  the  jiiilsc  «a\c  varies  aecordiiiir  to  the 
'i.:'..iil,\    of  the  walls  of  the  arteries.     To  iiliiicrstalid   why  this  should  he 

.,,,,    it    will    lie    well    for   a    moinenl    t nsiih'r    the    physical    eoiidiiioiis 

iipnii  \\liieli  the  iMiKe  \\a\e  di'pciids.  If  we  <Miiiiice|  a  piece  of  riirid 
i;ihe  with  the  inz/le  of  a  lartre  s\  lin-ie,  with  each  movement  of  the  liis- 
I. 11  a  wave  of  pressure  will  lie  transmitted  to  the  lliiid  in  the  tiilie,  aloiivr 

■  lii.-h  it  will  tra\el  at  siiidi  a  liiitli  \elocit\  that  it  will  arrive  at  the 
inc  end  of  the  tiilic  almost  iiistantaneoiisly,  and  iiicidcntally  the  out- 
law of  llnid  from  the  end  of  the  tiilie  with  each  com|)ressioii  of  the 
I'Miiip  will  ill'  cMic-tl.v  equal  to  that  represented  liy  the  iiiovemeiit  of  the 
j;>tun  If.  on  the  other  hand,  an  elastic  tlihe  is  employed,  it  will  lie 
inMiiil  that  the  sudden  increase  of  pressure  produced  liy  each  stroke  of 
'!'•  |iiiiiip  causes  a  distention  of  the  walls,  which  travels  aloiur  the  tiilie 
:is  :i  wave  a;  readily  mcasiiralile  \cloi-ily,  which  is  slower  the  more 
'  \t.iisilile  the  liilie.      .Moreover,   the   oiittlow    of   tluid    from   the   free  end 

!'  ilie  tiilie  will  continue  for  some  time  after  tlic  i-cssation  of  the  move- 
MM  111  of  the  pump.     What   happens  in  the  tiihe  with  each  dischai'Lrc*  of 

i  ■  lliiiil  is  that  the  |)ortioii  which  is  imiiicdiatcl.v  adjacent  to  the  pump 
i:  idcriroes  distention  aii'l,  heiiii;  elastic,  tends  immediately  afterward  to 
"•  -"il  ami  thus  exert  a  recoil  pressure  on  the  Iliiid  contained  in  the  tuhe. 

\-  .1  nsiilt.  i)resslire  ^\a\es  are  set  iiji  in  the  llnid  in  all  directions.  Those 
'i  ;i!  travel  hack  come  to  a  stop  liecause  of  the  iiistoii.  while  those  that 
!'  i\cl  distally  act  on  the  llnid  in  front  of  them  so  as  to  accelerate  it 
:  '  •!  liy  temporarily  raisiii<r  its  pressure  distend  the  next  sesrment  of  the 
'^-'1  wall,  until  the  end  of  the  tiilie  is  reached.     From  this  coiisidera- 

■  '1  t  is  clear  that  the  more  extensiiile  and  elaslic  the  wall  of  the  tuhe 
:^.  Ml'-  loiiL'cr  will  it  take  for  the  wave  of  pressure  to  travel  from  one 
'■  ■  1  to  the  other. 


'  >i 


LMH) 


I'lii:  (  iKi  I  i.A'i'iov   OF    rill.  iii.iMiii 


ri 


Alti'i'iit  inn  iij  till'  I'iili'  III'  ti'iiii>iiiisNiiiii  III'  til*'  |iiiIm'  uaxc  in  tlir  iirti'i 
ii's  ol'  iiiiiii  i|i'|>i'iiils  <'iitii'i'l\  iipnn  nil  ii|i|ilifjilii>ii  nl'  iIk-m'  |ii'iiici|ili'^ 
Wlicii  the  iii'tcricN  lit'i-iiiiic  liiiidt'iicii  in  nlil  a'.'*',  the  rate  itt'  tiaiisiniNsini 
111'  the  pulse  wave  i'^  iiiarkt'(ll\  iiicifast'ii.  The  iiiiisc  is  also  tl'aiisiiiitti'> 
iiinrt'  rapiillv  in  ilic  msscIn  nl'  tin'  Inwi'i'  cxlrciiiitii's  tliaii  in  (Inisc  of  1Im 
iippi'i",  siiHT  ill  till'  I'lii'tiH'r  iIh'  liliiiiil  M'ssi'ls  lire  snMicwIiat  iiiori'  riyiil 
i)t'lay  in  tlif  liaiiNiiiissinii  nl'  the  piiisr  \\a\«'  is  fui'lliiT  iiliscrvcd  as  him 
of  tlic  siiriiv  111'  .iiii'iiiiMii  in  .1  "ssi'l:  as  is  wrll  knmxn.  anrniisni  of  IIm 
siiliclax  ian  ailtiv  mi  nin'  siili-  caiisi's  a  di'lav  uf  tlic  pulse  nii  that  siil' 
flint   is  peiceplilile  t(i  tin'  Mlit'i'is 

The  Contour  of  the  Pulse  Curves 

l'"iir  niiiie  paitii'iilar  stinlx   ol'  the  I'Mlct  eonloiii'  of  the  pulse  wave,  ami 
espei'iallv    I'nr  iletiiniiniiiir  the  time  relationships  ol'  the  seeoiidary  wave- 


lu       -7  lli.ii^i.if:     ol      I    l..t;i\.  .i;i'>     .li..NiMi:;.i|,]t.        il.     tnl.i       t-T     itil  t  ..iJiM  I  I'M)     ml.,     tin-     liiliiiii     . 

.il<ry.    iuul    t.ii'Min    ;i    li.t.llr    -.1     \.irii.    whnli    ].,!--.-    ttii..ii:h    111.     (I.isii,     Ml.  nihr.inr    in    il- 

.iiiil    IH..\,,    l.\     ih,     iMi|..il-i     ..I    111,     1. 1 1    .iiiinl;    ..    ti.i.lii.ili.l    ...ill     li,i     ini.i-'jiiMK    llu'    i\'i 

'li.     ..-.  >ll;lt'..n'     ..t     III"    imkIIi'^ 


a  liirire  \aiietv  ol'  luitiioils  oT  vai-yinu'  iletriees  ol'  acelliiii'v  have  In  ■ 
elaliiirateil  for  eaeli  kind  of  ]nilse. 

Tiiose  devised   for  liieasuiilii:  the  /irissKn   jiiilsi    have  already   lieeii   'i 
M-rilieil  I  SI 'I'  pa;ri-  r_'T  i .  and  for  I  lii'  other  pulses  llii'V  are  as  follows : 

Vihicitii   I'lihi.      Mueh    iiiu'eiiuity    has    lieeii    displiiyed    in    the   elalioi 
lion   of  iiiethnils   for   rei-onliiii:  the   xcloeity   pulse.      Ill   one  of  these   r 

artei'.v    is   i-iit    ai'i'oss  and   tl lids   altai-li''d   to   a   tiilie.   into   the   iuin 

of  which  iirojii'ts  a  paddle  or  vane  arlieiilated  with  a  liirlit  lever,  win 
passes  throliudi  its  wall  i  see  ri".'.  .">7'.  The  \ane  floats  in  the  hi" 
stream,  and   the  outer  end   of  the   lexer  to   which   it    is  attached   is  c 

iiected  with  some  device  to  n rd   its  movements,  whieli  vary  with   i 

veloeily  of  liiiiodiiow  i  iicmodromoirraph  i.  Another  iiieliiod  eoiisisis 
the  application  of  the  instrument  known  as  I'itot's  fiilie  used  liy  pf 
ieists.     This  consists  of  a  hori/ontal   tiilie  liaxiii','  two  side  tiilies,  eaeli 


Till.    IIIiHiliM  IIW     l\     IMI.     Ml'lllll 


•-'tit 


ii'li  JH  ciinncflcil  at    its  ciiitct    riiil   wiili   a    iiiiiiiiitiK'ti'i'   aiiil    |iriiliiii'.'i'i| 
nlc  till'  liiiii/iiiitiil  till"',  \\lic:i'  tlic'v    ar-  licnt   at   ii|>|iiisili'  liu'lit  aiiifli-s. 


that    tilt'    iiiiii'i'  ('Mil    lit 


mil'   111'   tliciii     till'    |iiii\iiiial    tiilii'     pii'nits    up 


W 


t 


SD 


r-\\ 


il 


lll.IUI.int     to     <^•lv^      |.tttM  It'll      <'l      I'ltnt'^     mills     t-. 


;,     ,.ul-..       Im    I. ..11' 


Until    »tll    MM     lii'i.iii.i     ..I    l.itii.tl    i.n.-itr   .    Iitit    in    tin     |.i..Mni.il     i  li  II  h.inil  >     Inln     It     util 


v'li  1    111. in    in    llir    ili»lal.    In 


II    »ill    .-lUi.    Ill     .irtiili   I    lit     till     mIiiiIiv    III    Ho 


ll|;ii:r.itii    In    lllnstijli'    tin-    j.t  itii  iflr    nt     lAlmUUt's     I'liotn  lirtiMlol. 


iltontilit         'I'lii     Nil 


.'-    Iniilirr    tli.in    lli.il    111    /'    in     |ro|ioiiii.n    t..    tin      \i|.-.  ;lv      ■!     Hni     ..I     t'n     liliim;    ali 


J 

4 


ami  rccnrds  nut  iiiily  tlic  iattral   iui'nmiii'  Init   alsii  tin'  |ln'^^lll■l 
il    l>y    the    MHJili'ii    iiicii'aM'    in    Nclm-ilx     nl'    tin'    llnw.    wliiic    tin 


•JO  J 


Tin:  niiri  i.ation  ok  iiii;  iii.(h>i> 


^     I 


(ill'if     llic  ili>1iil  tiilx'     licintr  lu'iil  (liiwn  strcimi    n'<'<>rils  iin'toly  liitfi.,1 
|>r«'ssiiri'.     A  pliiitouiapliii'  rrcord  of  tlif  iiioNciiiriit   of  the  lliiid   in  tli> 
fuo  tiihf!*  K'vi's  till-  \rliicity   piilx-   ( M'l-   Fiir,  TiH).     Fur  {iliNNiolni^iir  p\ii 
posis  the  fiiiiii  (if  appaiatiiN  umiI  i»  ( oMstiMct*-'!  nx  slitiwn  in  Vi\t  TtO. 

I'lllpiililr  I'tilsi .  T'l  sti-mc  n  iffdicl  of  till'  piilpal)l(>  pllls(>,  tin-  m. 
••allt'd  splivuiimurapli  is  niiplovtil,  alllKHiu'li  a  tainl'inir  liavinir  a  liultm, 
in  tiio  ci'iilrf  uliirli  is  iiiadi'  In  press  on  tlir  arlciy  may  also  Iw  en, 
(iloyed.      The    eoiiiniuiiest    foirii    of    spliymnoirmph    is    that,    known    u- 

Pndu'eon's  (  I'itr.  (Kii.     It-eonsists  of  a  smail   Imlton  coin ted  vilh  ;i 

spfin;.',  tlic  niciM'inents  of  whiell  ai'e  transmitted  and  iiiairnitied  by  niean^ 
of  a  sxsteni  of  levels  eniineeted  vvilh  a  writing  point  arranu'eil  so  ;i> 
to  iiiseiilie   its  rnciveinents  on  a   nioxim.'  siirfaee. 


The  Analysis  of  the  Curve 

The   jieiM'ial   ennlonr  id"  liie   pnlse   wavis  taken   liy   any   ot   the   alioM 
methods  are  in  tieneia!  very  nnieh  the  same.     The  pi'essure  and  veloiiiv 


ii..,i  ..f  ii.. 


ritf,  'l.  Pill'**'  tr.irinir  <«["ln  |tnii>Kraru>  taktri  l.v  tpIiVKnuiyraph.  a  i/.  the  |> 
iiir\.:  '•,  till'  j.riniar)  ;  i".  llii-  ilu  rulit  »avi.  'I'li-u'  tii.irkr.l  in  liOti'*  i.l  .i  ^i ,  i 
(i,.|j    /'"■  </•■,'.  ..;v  I 


pulse  enrves  ate,  liowever.  not  nsinillv  taken  for  the  pnri>ose  of  olis.  i 
injr  the  eontonr  of  the  «a\e  hut  lalher'  for  uh'asnriiiir  the  difl'erenei 
pressure  or  veloeity  aetuali\  prodneeil  dnrinsr  eaeli  pnlse.  It  is  a  reii. 
of  the  palpalile  pulse  that  is  nsuail.\  employed  for  studyinf;  the  eonl,. 
of  the  «a\e  and  the  jiresenee  of  seeondai'y  waves.  A  leeord  of  tlh-  p. 
jialile  pulse  wave  i  l''ijr.  (il  i  shows  two  .separate  waves  on  the  deseeiiiJ!' 
linili  of  the  main  wave.  If  a  lai'U'e  mnnlier  of  similar  jmlse  enives  ::i 
taken    fioni    dilTerent    individuals    or    from    the    sanu>    individual    nii'!- 

dilTerent  enuditioiis.  it  will  he  found  that  of  these  two  waves  the  si 

(nn>  is  liy  far  the  nmre  constant:  and  if  the  waves  are  tii 1  in  relati 

ship  to  tlie  heart   sounds,   this  second   wave  always  « nis  iiiiinedi:i' 

after  the  SI Mil  sound,  allowance,  of  course,  heinu'  made   for  the  tii; 

l'ei|uired  for  the  jiulse  to  lie  t laiisniitted  from  the  lieart  to  the  ai' 
from  which  tiie  iiuise  tracini:  is  lieini;  taken.  If  the  olisei'vatim 
nunle  ver-y  earefnil.v.  it  can  indeed  he  shown  that  the  second  sound  ' 
responds  exactly  to  the  notch  which  precedes  this  v\  avo.     The  waves  ' 


1 


tm:   lil.<i<M)KI  nw    IN 


AKrf:Hlf>i 


2o:i 


I    |i>  lliis  iiiitrli  run   nut    lie   I'cliiti'il   tii  ilrliiiiti'  i'l>iiii','<'H  (ii'fiiri'inif  ill 

'''  hi'iiil.  K\  iih'iillx ,  tlii'ii.  Ilic  Mcniiiljiry  imlsf  wavfs  inr  imt  nil  nf 
.  iii;il  si'.'iiilifiiiicc.  Iiy  far  lln'  lii<»t  iiii|»'l'(jiiil  liciiiK  tiiii.  wliii'li  (icciirH 
iiniiniliiltcly  llt'liT  tin'  scr'dihl  mmiihI,  fjilli'il  tlic  iliiriilir  intrr  (<•),  llio 
I  rt.li  ill  fnnif  (if  it  lprjiii»  fiillfil  tlic  ilitmlic  nolrh.  Any  scciuiilai'y 
,;m<  occnniiiK  lici'inf  till'  iruTiilic  arc  callcil  i>riiliinilt<\  or  il'  tlicy 
•  ic'iif  (III  ilic  aMi-i-iiiJiiit;  liiiili  of  iIm'  itiaiii  piilsi'  \\a\<',  as  ihry  siimi'tiini's 
.I',  tliry  arc  call"''!  iiniurolir.  Wavrs  (iccuiiini;  alter  tlic  (licintic  arc 
r;ill(M|    fiiislltil  riilii'. 

Till'   ii'lativc   iiii|niilai nf  the  cljcinl  ic,  in   c(iiii|iai  isnii    with   Ilic   |irc- 

■  Im  inlic  ami  |)iislilicriitic  x\a\cs,  is  riirllicr  cvidcuccil  liy  Ilic  fact  that 
il  ;ilciiic  is  seen  on  a  sncallcil  In  iiutliiiniviiii).  which  is  the  tiaciiiu  oli- 
l:iiiiii|  liy  allouiiii.'  a  tiin'  stream  of  IiIimhI,  rsca|iin!,'  fmni  a  iiinlmli'  made 
in  ilic  wall  "iT  an  ailcry,  to  iiM|iiiiv;c  ni><>ii  a  ninxini;  >licct  nf  wliiti-  Mnt- 

tih/  iiapcr.    That  siicli  a  liaciii>;  >liii\\s  a  iliciipiic  liiit  mi  s iiidary  wave, 

iiMii.-atcs  that  only  the  I'driiici'  is  picsriit  in  the  Mocul  stream  itself,  ami 
lilt    tl thcr  SI mlai'.v    \va\es   nniNt    lie    iirmliieeil    liy   sunie   coiiilition 

<i      iir^   either    ill    the   clastic    lisMlc   of    ihe    walls   of    llie    lilnnil    \e>sels,    or 

ill  tlie  clastic  i>roii(itics  of  the  instiimicnls  iisi'il  for  taking  the  pulse 
li.i-iiii;. 

The  Dicrotic  Wave.  I'eeMiise  of  its  oli\  imisly  u'lcatcr  siuMiidcancc,  \\c 
-!  1  liist  of  all  cipiisicler  the  exact  cause  of  the  (licintic  wave  ami  of  the 
i  "nIi  pieeeiliiiir  it.  Thcorclieally,  I  \\  o  po>.-,ilp|e  causes  iiiiirlit  explain 
ill  »\a\e:  either  il  is  line  to  sonic  >eiiinilai\  wave  set  up  at  Ihe  heart, 
!t  is  ilepemlcllt  upon  w.ives  lelleelril  from  the  pel'ipheiy  of  the  cir- 
i'i!:itinii  liacl\  aloiii/  the  liloml  streaiii,  just  as  seeomlary  wa\cs  aic  re- 
''  •'■■■I  iVoiii  the  w.ills  of  a  lull  of  watci'  when  a  stone  is  thrown  in  the 
11 'il.  1-  III  eoiisideiiinr  this  sceoiid  possiliility,  we  are  of  course  maUinsr 
ill    aNviiiii|)tion  that  at   the  I'lids     I    Ihe  iiiterial  system  there  is  ;i  sudden 

1' ^!»t.iiiee  to  the  onward   moMinelil    ipf  M 1.     The    freiiuent,  hraiichins; 

'  '  ill  cn'ciifs  when  the  arleiioles  open  into  the  e;ipillarics  no  doiilit  of- 
iiiiiiiy  opportunities  for  the  lelleetioii  of  |)uli-.  waxes  hack  to  the 
I  'I  I'lit  ihcsc  waves  must  he  relleeled  at  sii -h  Naiyiiiu'  distances  alon;; 
''■■  .  t'Tial  system  that  there  can  he  little  oppoiluiiily  for  them  to  lic- 
.iililed  to<;ctlier  so  as  to  form  a  wjive  of  siiflieieiit  majinitudc  to 
n  ,1'  iivijf  perceptilile  in  tlie  hlood  Mowing'  i'l  Ihe  larirer  arteries.  These 
■  .1  -  ale  relatively  so  small  and  they  oceiir  .'it  such  dilVerciit  limes  thai 
)!•■  lilt  result  of  their  a<Idilion.  so  far  as  the  produetion  of  ii  larirer 
V  •■  is  eoiiecrncil,  must  he  praclieally  nil.  Not w  ilhstaiidinir  these  con- 
s;i  r:itiiiiis,  It  IS  ]>ossiliie  that  under  soiin'  eomiitioiis,  sm'li  as  in  cases 
it'  'i'.'li  aiteiial  tension,  certain  of  the  prcdicrotic  or  postdicrotic  waves 
111.,;.  Ill   due  to  the  above  causes. 


i  \ 


i 

t 

I 


I 


2(»4 


riii;  (  ii{(  ri.ATKiN  ok  tiii;  iii.oin) 


'I'liMl   //"    ilirriillr  Is  nut  n  lulUclnl  irari    is  ck'iiily  c'staltlished  by  tli 
I'in-t  tliiit   if  tlic  (listaiit,    licluccii  llic  ilicrutic  wavf  iiiul  the  main  piil- 
wave  is  iin  asiin'd  at  (lilVcrcnt    |mints  ol'  tlir  aitorial  stirain.  it   will   .li 
wavs  hv   riiiliiil  til  111'  tlif  same,  wliicll  olivioiislv   would   iint    lit'  tlif  ca^ 
were  till'  difiniic  wave  ri'lU'cU'il.     11",  U<f  v\in\\\)\v.  \\v  wi-ve  to  oxaiiiii 
the  coiiloiir  tif  till'   wavo  prodtu'cd   liy   tliniwiii^   a  stone  into  a   tiili  <•■ 
uatiT.  we  sIkuiIiI  lind  that   near  ll.c  ed'^c  tlu'  secondary  wave  was  vci 
cluM'   1(1   tlic    main    wave,    \\liei('as   ni'ar   the   renter   the   secondary    wa\. 
would  iiccui'  iMUi'li  lal<'r. 

Our  iirolilcni  tln'iet'ori'  iiaiiuws  itself  ilown  to  an  investitration  ■ 
the  cause  fur  the  dicrotic  wavi'  at  llic  central  end  of  the  cireidation.  I 
occurs,  as  we  lia\e  seen,  innncdiately  after  the  lieiiinniuH;  of  diastol' 
That  it  can  not  lie  due  to  anytliins.'  takini;  place  in  the  ventricle  itself  i~ 
evidenced  liy  tlic  fact  that  siu-h  a  wave  is  alisent  from  an  intracardi. 
|ircssurc  cur\"  i  sec  paye  |."ip.  ;iltliouj,di  it  is  present  in  the  very  lieuii 
nin-r  of  the  aorta.     Now.  the  only  structures  existing  between  those  t\\- 

points  which  could   In-  held   i  es| silile  for  this  wave  arc  the  sciiiilu!!.;' 

valves  a  conclusion  which  is  sustained  by  the  fact  that,  if  the  aoit;- 
\alvcs  are  rendered  incoinpctent  by  hookiiiir  them  back,  or  if  the  i>nl-' 
beat  is  examined  in  patients  sulVerinjj:  from  an  aortic  insuflieieiu-y.  ii 
will  be  I'ound  that  the  dicrotii-  wave  is  not  lu'arly  so  evident  as  usual. 

To  iinderstMnd  lioir  Iht  niln  s  iin  n  iixnisihh  fur  Ihi  praiJuiiitiii  of  ^• 
iriifi.  the  mechanical  clianires  occurrin>.r  at  tin'  root  of  the  aorta  mu  ' 
be  ili,irl\  nnderstood  i  sc  pa'^'c  bVi  i .  The  slretchiny:  of  the  ela.stic  «,i!!' 
of  the  aorta    whicli   occurs   with   each   systolic   outrush   of  blood   is    l'-' 

lowed    by    a    iiowcrful    and    suddei ntraetion    of   the    stretched    wal  - 

;ind  the  pressure  thus  brouuht  to  beai'  on  the  column  cf  blood  in  tin i'. 

tends  to  impel  it  both   forward  and  backwai'd.     The  forward  umvei ' 

adds  itself  to  the  wave  of  increased   pressure  already  produced  by   i' 
\en1rieular  contraction.     The  backward  <'omi)oneiit  travels  as  far  as  •    ■ 
semilunar  \al\i.  from  which  it   is  rcllected,  ami  now    iiniceeds  periplp 
ally  alonu  the  blood  stream  durini,'  the  tinu'  at   which  the  'iri<:inal  pi 
>ure   )iulse    is  ijeclinin!;.      It    therefore    imjiriuts   itself   on    the   pulse   tr 
iiiLT  as  a   separate   wa\e,  and  does  so  all   the   nmre  markedly  when   '■ 
decline  in  thi'  main  pulse  wave  is  rapid,  as  in  eases  in  whieli  tile  peril" 
eral   resistance   is   low.   but    fails  to   be   prominent    when,   on   aceouni 
a  hi'.'h  peripheral  resistance,  the  decline  in  the  main  pldse  wave  is  lai 
This   explanatinn    coineides   exactly    with    the    well  known    cliiucal    ; 
that   the  dicrotic   wa\e   is  conspicuous   in   pulses  of  low   tension,  bui 
markeij  or  absent    in   pulses  of  hij;li   tension. 

One    ]>oint    remains    to    be    consKJered,    ami    that    is    tin    nnisi     hit- 
siiihii  II  ihrlliu    III  tin    iiiiini   iniii    at   tl -ssation  of  the  \entriculai 


I'lii:  lu.uoDKi.dw   i\  riii:  AiiTnm:; 


HO.') 


|Mit.  Idi'.  it  iiiisjlit  lie  siiid,  why  sliniild  llicre  lie  siidi  a  smMfii  fall  in 
|.r.ssiiif  iiciir  llic  licarf,  wlicrcas  toward  tlio  j)crij)li(iy,  as  we  have  socti. 
ihf  pit'ssiiro  liotwccn  tlio  licartln-als  tends  Ui  In'  inaiiilaiiicd  on  account 
Ml  till'  clastic  recoil  of  tlie  stretched  ai'teria!  walls.  The  explanaticm 
ii^iiallv  (liven  is  that  the  sudden  cessation  of  o\itllow  of  Mood  fi'oni  the 
.iiliicle  at  the  end  of  the  s])livi:niie  pei'iod  causes  a  nes^ative  i>i'essure 
In  lie  inodnced  in  the  lilood  at  the  hcjiinnin^  of  the  aorta,  thus  tendiud 
Im  c.iusc  a  relhix  of  hlood  touaids  the  heart,  the  en'c<'t  of  which  is  (  1)  to 
I, nine  the  closed  valves,  and  ci  to  pidduce  the  reflected  dicroti<-  wave. 
1 1,  wiiiie  iluid  is  llowinn  mulcr  pressure  ahniir  a  tulie.  the  llow  is  sud- 
il'  iilv  arrested  by  tnrnin<r  a  stopcock,  it  is  possilile  liy  the  use  of  inaiKini- 
( t<  Is  to  show  that  a  nefjative  wave  is  set  up  iuinndiatelv  licyond  the 
si.ipcock.  and  that  this  neirative  \\a\f  trascls  alony:  the  tulie  at  a  rate 
.|.|iiiidinu;  on  the  elasticity  i>f  its  walls. 

Causes  for  Disappearance  of  the  Pulse  in  the  Veins 

III'  ilisa])i)earaMcc  of  the  pulse   in   the  capillaries  an<l    its  eiiiiscq\ient 

, '-I  !iri'  In  the  veins  \\c  have  ah'catlx    seen  to  In'  owiuir  to  tin inhincd 

•■■  of  the  elasticity  of  the   \  csscl   walls  and  the  i)erii)hcral    i-esist- 
account    of  these   two    factors   the    pressiu'e   conveyed    to   the 

i,i I  ilniiii";  systole  is  stored   up  to  he  ielease<l  durin<r  diastole  liy   ihi' 

ii.nil    uT   the    strelehcil    vessels.      Sonn-tinies,    howcM'r,    the    pulse    jicls 

ihi'iiLili  to  the  M'ins,  either  liecailse  the  elasticity  of  the  vessels  is  not  so 

iii.ii  l;i'i|,   or  hecause   the   peripheral    resistance    has    lieen    lowered    i\aso- 

.liiii.itidii   .     In   iiaticnts  with   hardened  arteries,  or  in   normal    indi\idu- 

.<.\-    iller  taUiiifr  nitrite,   which   dilates  the  peripheral   ai'teiiole^,  a   l)ulse 

n  .1}   ccinie  tiiron^ifh  at  the  ])erii)hery  and  appear  in  the  \cmis.     This  nni.v 

'    .  ailed  the  pei'ipheral   venous  )iulse,  and   it    is  to   lie  carefully   distin- 

^hed   from   the  central   venous   pulse  oliscrvcd    in    the   larnc   \eins.   as 

::•  'hr  root  of  tlic  ucck,  licforc  any  vahcs  ha\e  inlervi'iied  to  hlock  the 

'      '^mission     of  the  auricular  pressure  waxc  hack   iiito   the  cohunn   ot 

'       'I    in    the   veins.      If   a    inilse    i>   seen    in    a    larye    \i'in    ami    there    is 

■1    ilit  as  to  whether  it    is  peripheral  or  central   in  origin,  this  doulit   can 

iinnn'iliately   renniveil   liy    locally   const  rictiuL'   the   vein;    if   ihi'   jiulsc 

;  'lipheral,  it  will  disapjiear  on  tin-  heart   siile  of  tli mstriction;  if 

-  central,  on   the  side  awa\'    froni   the  heart. 


h^-u 


CIIAI'TKR  XXIV 

TIIK    f{.\TH  Oh'   MOVKMKNT   OV  TIIH   HI.CHM)    IN   Till': 
I'.LOOI)  VKS8KLS 

Siiic<'  tlic  nlijcct   iif  tlif  ciiciilMrmM  is  lo  iiiiiiiitiiin  iui  iultMniafo  iikim 
iiu'iit  111'  lildiiil   ill   tlic  tissues  jiikI  c-iiiillnries,  it    is  cvidfiit   tli;it   licsidi - 
iiit'.-isuriiiir  the  iircssiiic  nl'  l)l(in(|l!(p\v.   \\c  slumld   also  nicasurc  the   r;ii 
111'  its  iiKivi'iiit'iit.  Ill',  a.s  it  is  iit'tcii  cailiMl,  tlic  mean  velcieity.     Tiiis  ineasiii' 
iiient   may   lie   uii<1ertakcii   eitlier  for  a    ".'iveii   vessel    or   for   a   eoniplii' 
vaseiilar  area,  siieli,   for  exaiiipie.  as   tliat    iif  one  of  the  viseera   or  oim 
of   tlie   extieiiiiiies     tlie   mass   mo\ cmeiif   of   llie   lilood.      Or   instead    I'l 
iiieasiiriiiir  liie  mean  \(|oeil\    we  may  desii'e  to  know    liow  lonjr  it    taiv - 
fur  a  particle  of  lilood  to  traverse  a  iriven  vascular  ai'ca.     Such  a  me:i^ 
uiciiient  is  called  tlie  rlrriiliilidii  Ihiif;  it  do(>s  not  at  all  tell  us  lio\v  lim- 
it   takes   for  nil  the  Mood   to  jiass  lliroii^di   the  "jiven   area,  Iiut   only.  ,^ 

slati-d.   the    time    ici|uirc>d    for  the   circiiliition    of  a    fr,-iction    of   the   M I 

throiiudi  a   particular  Held. 


I      1 


VELOCITY  OF  FLOW  IN  A  VESSEL 

Speei.il  iiiclliods  lia\e  licell  de\  isecl  for  the  measurement  of  each  ' 
these  three  \<locities.  I'or  the  measurement  of  the  velocit.v  of  ll" 
thi'oUirh   a    main   artery   or  vein,   methoils  similar   to   those  em])lo.\'ed   I' 

li.\draulic  eiiiiineeis  are  emplo.vcil;  th;it   is  to  sa.v,  the  volume  of  M 

in    culiii-    <'entiiiictcis,    which    passes    a    iriven    i>oinf    is    meas\u'ed    for 
iiixeii   time,  and   'he   I'csiill    dixided   li\    the  cross  section   of  the   vessel 
the  point   of  oliscrx  alinii.     Tlie  result   j;i\i's  us  the  iiiku)  IIiikiI  vihui'' 
To   measure  the  oiitllow    of  lilood   ill   a    i;i\('U   time,  the  simjilest    liietli 
would    lie    to   cut    .icioss    the    Vessel    and    colh'ct    till'    Mood    ill    a    SII''"'"'' ' 
liut    oli\  ioiisl.\-    in    this    nntliiid    an    error    would    he    introduced,    licciii: 
eiittini.'  the  vessel  would  lower  the  peripheral   rcsist.'iUee  and   remo\e  i 
natural  olistructinn  to  the  tlow  )iresenf   in  the  intact  animal.     Moien\' 

till'  heiiioiiha'je  would  in  itself  iiit  riiduce  a  ilist  urliiii!,'  tactor  on  a w 

of  the  loss  of  eirculatiii!,'  tlilid. 

To  make  silch  iiieasuremeiits  of  aii.\  value,  it  is  i)h\iously  necessarx 
retain  the  |ieiiplicr,il  resislaiice.  l''or  smaller  vessels  this  can  lie  il' 
liy  introdiicintr  in  the  coiiise  of  the  artery  a  Imii,'  v'lass  tulie  lieiit   in  i 

2eC 


RATE  OF    MOVEMKNT   OP   THE   III.OOD 


207 


sliiipc  of  llic  lollor  V  (V\<i.  (VI),  or  Ity  iiuM't-ly  allowing  the  vessel  to 
I.Iiim!  into  a  {rrndnati-d  tiihe  and  seeing;  liow  long  the  blood  eolunin  takes 
|o  travel  from  one  end  to  the  other.  This  method  is  of  eonsidcrablc 
viilue  in  measuring  the  veloeity  of  How  from  small  vessels  sueh  as  the 
viins  coming  from  glands  and  nniseles.  For  larger  vessels  a  so-ealled 
stniwKhr  is  emidoved.  There  are  numerous  forms  of  stromuhr;  that 
sliown  in  the  diagram  (I-udwig's)  (l-'ig.  (i'i)  consists  of  two  glass  bulbs 
united  above,  and  eonneeted  below  with  lubes  that  open  Hush  with  the 
Mirlaee  of  a  brass  dise.  This  is  i)ivote(l  at  its  center  with  another  similar 
|iliitl'orm  also  having  (lush  with  the  surface  the  openings  of  two  tubes  eon- 
11. (ted  with  the  cut  ends  of  tlic  artery  or  vein.  In  a  certain  position  of 
Hi.'  platform,  the  tubes  from  the  artery  or  vein  are  exactly  opposite 
tlhiM'  of  the  bulbs,  so  that  the  blood  ran  How  from  one  end  of  the  vessel 


;.    9 

ill 

f    1 

|-« 

i^M 

■' 

\1 

'    3 

J 

s 

•  } 

■1 

III 

l'Mrin«   of    ,ip,.ar.ilii^    f'.r    mk  ;i<iir<  tin  iit    i.t'    ti!..i.il    iilm-iliis. 

■,'»    li,ii;  ..i(.,i.,.,rml.  ■.       'I'I.,-    I.I 1     11--. I    i-    .itt;i.lii.l    t.      ill.-    Iv.ii    ^li.rt    ^i.li-    lul..s. 

.,•   ■     ,  1,.    Ill,-    |..,..ili..M    ,f    th,-    -.I..!"...  k.    ihi-    I.I I    ll..»-    mini     .liT..tly    lr,iw,.ii    lluiii    i.r 

llii'-ijli  l!.    I    -li.,i-,-l   kI;i-^   till..-. 

■  I. :i  ii.it'-  ^li..iinilir.  Til.-  liil.,-  .-11  111.  l.-vv.r  .n.l  ..f"  ,  ^i,  li  ..f  111.-  Iw-  >;l:i-s  I'ull.s  |ii.-r,.- 
.ir,,,!,,r    I..,,,    i,|;it|.,nii    .1.1.1    (11.1    lli.-h    VMlh    il-    s.iil.i.i-.       I'lii-    pLill-iiin    pi.-.T-    ..1    il~    i  i-ilt.  r    ,.n 

^■niilir    l-.w.r    |il.i!f..!iii     wilh    tw..    ..|.(  iiiiik;-.    i.iMti..I..l     witli    tin-    tiil.c*    thai     l.a-1     to    tlie    l-l 1 

-•;]. 

tdiiuli  the  bulbs  to  the  other  end.  To  use  the  instrument  the  proxi- 
al  bulb  is  lillcil  with  oil  ami  the  ])cripliei'al  one  with  ithysidlogic  saline. 

ic  (lip  is  tlicn  removcil  frum  tlu iitral  enil  of  the  artery,  anil  the  blt)od 

•■.\s  in  and  displaces  the  oil,  which  in  turn  ilisplaecs  the  saline  in  the 
'l|ilifral  end  of  the  artery.  When  the  bloixl  has  risen  to  a  mark  on 
'■  tube  .iiiiniiig  the  two  bulbs,  the  inslrunicnt  is  rapiilly  rotated  so  tluit 
'  nil  is  brought  back  a-rain  into  the  proximal  i)ositiiin.  the  ititation 
'm,'  elTected  so  (piickly  that  there  is  no  ilistinct  iiiterru])tion  in  blood- 

"■.    The  i)])cr;ition  is  lepeatctl  in  this  way  for  a  given  pcrittil  of  time. 

inting  accurately  the  number  of  revolutions,  then  nmltiplying  the 
.'iilicr  of  revolutions  by  the  capacity  of  tlie   bulbs,  we   get   in  cubic 


I   I- 


I  •! 


-  !/ 


1  • 

i 


¥ 

V 


■JdS 


Till:  rii!(  ri.x'niiN  ni'    iiii;  i  '  ihid 


(■(■ntiincti'rs  tlic  imiiuiiit  oT  lilood  lli.it  liiis  llow.'d  tlirmitrli  tlie  iiistniiiifi 
ill  ji  <li-liiiilf  unit  of  liiiii'.  Tliis  -rivt's  tis  tlic  Vdlimif  lli>w  iiiul,  if  ll 
icsiilt  is  (liviili'd  liy  till'  iMoss  sci-tioii  of  tlic  \ csm'I  ill  siiiiiiic  cfiit imct.'i 
we  oliliiiii  wliiit  is  kiiciwii  iis  tlic  iiicjiii  lineal  \cl(M-il.\-.  .Many  iiioditii'. 
tioiis  have  Keen  made  of  tliis  iiisl  iiiiiienl.  Init  it  is  iinneeessaiy  In  <;<>  im 
llielii   lieie. 

'riie  uclicial  result  of  siieii  iiicasiirciiients  lias  liceil  fn  slinw  tlial  ll' 
lineal  xcliieil.v  is  iiinrsihl  iniiiinrliiiiKil  In  llii  i  nis.i  sictidii  of  the  vesM 
at  the  i>oint  of  oliscrx  alioii.  It  is  ol)\  ions  that  the  voliiine  of  lihu. 
llowiiii;  out  of  tlie  heart  to  tin'  aorta  in  a  iriveii  time  is  exactly  e<|U, 
to  that  tiowiii^  into  it  liy  the  vena  cava,  and  likewise  that  the  voliin, 
ilowiiiir  into  an  or<iaii  is  csactiy  ei|iial  to  that  wliieli  liows  out.  (.'on- 
i|ilcntly  the  lineal  velocity  will  lie  inversely  proportional  to  tlie  se. 
tioiial  area  of  the  vessel.  The  principle  is  the  same  as  that  which  yo 
ems  the  \cloi-ily  of  How  of  a  stream:  when  the  lied  is  narrow,  tin'  en 
rent  is  swift,  Imi  it  liecomes  slnjiiiisli  when  the  lied  is  wide.  If  ll 
arteries  were  .if  the  same  ealilicr  as  llic  \cins,  the  mean  velocity  of  t! 
Iiloodllow  tliroiiuli  the  two  would  lie  the  same,  hut  actually  it  is  iini'  : 
•ii-catcr  in  the  arteries  liei'ails.'  the  liinien  of  these  at  a  irivcii  iioint  in  ll 
eireiilation  is  oi.l\  from  oiie-third  to  oiiedialf  that  of  tlic  eorrcspoiidii 
\  ein. 

h    must    Im-    niideistiMMl    that    wc    are    tlealinii    ahove    with    the    me.- 
velocity  ill  a  iihil  of  time,  and  that  there  must  lie  coiisidcralilc  altcrati- 
ui;li  raeli  s,\sioh-  and  diaslnl,-.  emisl  itntinn'  the  velocity   pulse   (  paye  "Jm 
'{'he   dcyi-(-c   of  this  alteration    with   eacli    velocity   pulse   is  liiilch   less 
the  periplierx    of  the  eireiilation  than   near  the  heart.     As  the  periple 
is   reached,   the    llow    heeomes    more    uniform.      It    must    further    he 
mciiihercd    that,    altlKiliuh    the    mi'aii    velocity    depends    esselltiallv     up 
the  area  of  the  vasculai-  lied    .vet   it   is  suli.jeel   to  eoiisidcralile  variatio   - 
as  a    result   of  chaimes  eitlicr   in   tin-    force   or   rate   of   the   heartlicat 
in  the  t'aeility  of  oiiltlow    from   the  ends  of  the  arterial   system     that 
i-haiiues   in   periphiral    resistance. 

It   is  iisiiallv    stated  thai   the  mean  lineal   vclocit.v  in  tlic  carotid  art' 
is  alioni  :'.iiii  niiHiiiieters  pi-r  sei-niid;  and  in  the  .iuu'ular  vein,  alioiit    i     ' 
millimeters;    w  h(-ieas    in    tin-    i-apillaries,    where    the    total    area    of 

vascular  lied  has  1 me  enornionsly   iin'reasccj,  lieiiiy;  perlia|is  sonii'   ~     ' 

times  that   of  the  aorta,  the   velocity  of  tlow  is  only  aliout    half  a   in 
liieti'r  tier  s md 

MASS  MOVEMENT  OF  THE  BLOOD  IN  A  VASCULAR  ARE/ 

Methods.       In   consiilel  inu    lln    lilndilllulf  nr  iiKisa   iiiori  i)l(  III   III    llii    ll        ' 

in    the   ditTcreiit    reiiions   of   the    liody.    it    is   usually    more    jiraeiicai     " 


k 


R\Ti-,  OK  Mftvr.MrNT  OP  Till:  v.i.fxm 


20t) 


iii.-asuro,  1U.1  tlio  nioiin  lin.-al  vclccily  of  tin-  iiillow  in-r  iind  niitllowiii}? 
1,1.M)(1,  but  ratli.T  Ik.w  iiiiiiiy  (Mil>ic  .•cMtiini't.'rs  ..I'  1i1"'m1  aic  travcisiiii; 
ilio  part  per  KM)  j:iaiiis  of  (ir<ran  or  tissue  per  niiil  ol'  time  Siicli  iiicas- 
arrmciifs  may  he  ma.U-  in  a  vavioty  of  ways.  If  tlicri-  ar«'  l.iit  oiio  art(>ry 
Mi.l  0111-  vein  to  tin-  part,  tin-  sti-omiilir  may  of  cotiisc  hv  (•mi)loy.>«l.  and 
It  may  be  ins.Mtc.l  in  .'itlicr  the  aitciial  or  tlic  venous  eiieiiit.  Ftu' 
iiieasuniifr  the  mass  movement  of  tiloo.l  tlironirli  sueli  lar-re  visepia  as 
ihe   liver,   this   is   indeed   tlie   only    nietiiod    tliat    ean    be   employed,   the 

,;ronndir  heinir  inserted  eitlier  in  tli mrse  of  the  (H.rtal   vein  and  he- 

|,;itie  arteries,  or.  better  still,  in  the  v.-na  .-ava  .jnst  below  the  oiienintrs 
.,1'  the  hepatie  vein,  the  vena  cava  beinu:  slnit  otV  for  a  moment  between 
liie  liver  and  the  h.-art  and  tln>  blood,  as  it  Hows  from  the  heiiatic  vein. 
■illowed  to  eolleet  in  the  stnniudir.  For  other  orirans  an.l  tissues,  how- 
,',.r,  methods  which  do  not  involve  any  interf.-rence  with  the  blood 
.  -scis  may  be  emi.b.yed.  One  of  these  is  the  so-ealbd  /»/<,'/< .i/.s»H»/rr(///»i<' 
„„ilii,(l  of  nrnilir.  An  orv'an.  such  as  the  kidney,  is  enclosed  in  a  plethys- 
•i.i..'1'aph  (w'e  pau'e  2-»l)  and  while  :i  reeoi.i  of  its  v,.lunic  is  bcin^' 
i.^.-ribed  on  a  (|uickly  revolvinir  drum,  the  vein  is  suddenly  clamped. 
uiil,  the  result  that  the  kidney  volume  expamls  in  i>roportion  to  tho 
„;,,.s  of  blood  flowinjr  into  it.  ^Vh<•n  the  expansion  has  reached  a  cer- 
t;„n  de-rree.  the  clamp  is  removed  and  the  bloodflow  allowed  to  pur- 
-,,  :.  its  course.  It  is  then  an  easy  matter,  by  jrraduatinir  the  plethys- 
!::^-rai)h,  to  determine  how  many  cubic  c.-ntimeters  of  blood  must  havo 
!'  rd  into  the  oru'an  in  the  uMven  time.  To  avoid  serious  local  asphyxia 
r     !ic  tissue,  the  clamp  must  be  api)lied  to  the  vein  for  oidy  th-  briefest 

'  .      MJ  of  time.     This  meth.xl  may  als,.  1 mpb'yd   for  nu'asurinsj  the 

,lilou    through  the  extremities.     Thus,  if  the  arm  is  enclosed  in  the 
!       iNMUo-rraph    (Fiji,    ti:?'    and    a    ban.l    enei  ■•'■nu;    the    arm    above    the 

-mo-iraph  is  ti-rhteiu'd  so  as  t nstrict    ,        veins  but   not  the  ar- 

'he  rate  at  which  the  volume  of  the  arn\  within  the  i)le1hysmo'irai>h 
will  corresp.m.l   to  the   rat.>   at   which   bloo.l    is   llowine-  into  it 

I'.i  the  purpos.'  of  TMeasnrint:  bloo.l  How  throudi  the  upper  or  lower 
.\!  .milics,  II  mm-h  nnire  serviceable  rllninil  ni>  Ihod  is  that  of  (i.  N 
-.     art.     This  .lepends  on  the  priuciitle  that,  provided  the  blood  passin- 

11,   the   thinax    to   the   hamls  or    feet    is   of   ( stani    temperature,   the 

!.•  .11  which  heat  is  dissipated  from  the  hau.ls  or  feet  will  be  directly 
■ipnitional  to  the  rate  of  movement  of  the  bloo.l  throu-h  thes.-  parts, 
.itunately  f(U-  the  method,  the  hands  particularly,  but  also  the  feet, 
V  MK.rc  nV  less  perfect  radiators  at  least  they  arc  to  this  extent,  that 
the  temi)erature  in  their  environment  is  not  much  bi\»<'r  than  the 
•niperature  of  the  blood,  then  while  this  is  traversiuir  the  part,  if   will 


I  * 


L'Kt 


Tin:   riKcl  I.ATIKN    (iK    'I'm:    lll.duii 


!f  5  ■ 


IciM'   llt'Jit    to   tllc   ('M\  ildliliii'lll    until    till"   iillttlnw  iiiir   |,|.   \-,. mills    Mdoil    is   :i 

cXMi'tiy  till'  sMiiif  t(iii|itiiitiii-i'  jis  tli(>  ciix  innimcht;  I'm-  cmiiiii)!!',  if  lii. 
li.iiid  is  pl.icul  ill  \v;itfi'  lliiit  is  a  little  ciioltT  tliiiii  tliat  ot"  tile  IpIikmI, 
iiiiil  till'  tfiiiiiriiitilli'  iif  til"  IiIcmkI  in  ciiic  ol'  llic  liiryc  veins  ol'  the  liaihl 
is  iiie.isiireil,  it  will  he  lnuiicl  to  lie  tlie  same  as  that  of  the  water  in  tin 
water-Iiatli. 

To  nieasiiie  the  rate  of  Ijnw.  theii'foic,  we  niiist  aseertaili:  (1)  ho\'. 
iiiiieh  heat  has  l.eeii  i;i\cn  out  hy  the  part  to  the  water  siirroiindinu'  ii 
ill  a  trixeii  lime,  ami  r2  '  the  ililTeienee  in  temperature  of  the  inllowinL' 
(arterial  I   ami  oiitllowiiii:  (veiioiisi    lilooij.      We   iiieasiiir  the  amount   <•! 


"^-^  r"~ 


I','-'  '''  I'l'llu-M  ■.i.-'.,|.li  fn,  ,,,n,,!,„i.  V. .1,111,,  ,}':r<i:,^  v:  l\„  \v.,u.\  jni  1  f,,i,;,,ii,.  llv  ..1.. 
."L..  ,„■  r.,u.  „,,l,  „!.„!,  .1,,.  v..lmii,-  „.,■,.,-,-  „1„.„  ,1„  .niu  i.  ,  .„n|„ , --,  ,|.  ,|„  i„  ,„  i„„>;  „„,, 
""     ■■' I    '■""    I'l'    'l.l,i„ 1.       i|-,,,„l    J.i.lv.c.i.   I 


heal   -iiven  out    to  the   water  in   I'aloiies.  a  ealorie  heinu'  the  amount 
heat    re.|uirerl    to   raisi'    the   teiiii)eratiire   of    1    e.e.    of   water    from    (I      ' 
'"    '      •'•      Suppose,    f,,i'   exjimi.le,    ;i    ii.-nid    were    placed    in    :!,(l()(l   e.e, 
"■■'''■''  '■''   •'•'     *'■•  "lid   that   afli'r  ten   minutes  the  temperature   had    ri^, 
to  :!.•!,:,     C.  then  the  amount  of  calories  driven  out   would  he  :!,(l(i()  x  (I .") 

'■"'""•      -"^iii alnri.'s   e.pud    culiie    centimeters   multiplied    l.y   ehaiiue 

lemper.-itiire,  it  follows  that  if  we  divid.^  the  lisrure  repres.'iit'iiijr  them  ' 
the  ii-liuilhi  ohserved   dilTerciice   in   temiieratlire  lietwccn    iiiflowin-;   a 
outlh.winir  l.!....d.   the   result    .unsl    eipial   th(^   lilii,.!.er  of  euhic   ..riitilMiI. 
of  IiIo.mI  that  lias  llowcd  thniii-h   the  (.art.     The  temperature  of  the   ■ 
llowiip.'  Mood  li;is  heeii   found  to  he  pijiet  ieally  identical   with  tliat   of  t 


RVIi;   (iK    MiiSIMI  NT  OK   'llli;    IllJHH) 


ill 


IIImII 


111  iiml.T  till-  hiiiUMi.-:  w  Ikmc.ms  i.l'  im.iii-  •  tin-  t(iii|i''iii1iii('  nf  llic  vciiiiiis 


s  iilrciHly  cNiiliiiiiiMl,   is  ciiniil   to  tlic  iiiciiii   1ciii|H'rii1\iri'  "f 
\,.itir  (lurin-r  tlic  lilin'  lli;il   tin'  liMinl   wiis  iiiiiiuTsi 


tliP 


il  ill  it.     Fiirllicr  do- 


.r  llu'  Icclmic  (if  this  iiii'tlin.l  will  !»■  foiiinl  I'iscw  Imti-.  Uiit  it  iiiay  h< 


.1  licit'  tliMt    it    is  cxtrciiM'ly   siiiipli'  iiini  iH'clll 


lie.  ;in.l  th.-it    it    rciiiurcs 


liMiiiiii;.'  iiinn'  tliiiii  (  Il  nil  :i<Tiiiiit<'  tli.iiiionictcr  iiinKins  lictwocii  i\\'» 
III    ('.  and  r>(l    ('..  witli  ii  scili'  so  (Innvii  mit  tliiit  i  (•a.Iiiij.'s  i-!iii  lie  inndc 

,,,  I, r  a  d.".Mt'(<.  and   fJ'    a    veil  .tusI  i  in-tcd   vessel    «<  alimit    :!.l"t() 

rr  capacity,  witli  dmililc  walls,  the  space  l.ctuccii  tliciii  hc'i.u'  jiai  U'd 
\  iili  sdiiic  heat  iiisiilatiiiir  material  such  as  y:iniiiid  cnck. 

Results.  K'eirardiiiv'  tlie  icsiilts  ohtained  with  these  iiiethnds.  it  lias 
hull  rmind  that  tlie  hlund  supply  for  eacli  ll'O  -jiaiiis  of  tissue  in  the 
\  iM-i  ra.  as  mcasiired  hy  the  si  rniiiiil'-  iiietliud.  is  almiit  as  rnlhiws:  stmiiach. 
■Jl  ic.  :  intestine,  71  ec.;  spleen.  ."iS  c  e.  ;  liver,  arterial.  'J")  e.e.  ;  liver. 
v.iK.ils,  Tilt  c.e.  ;  liver,  tutal.  S4  e.e.:  l.iaiii.  I:'.ll  e.c.  ;  kidney.  I.'.O  e.e.;  thy 
i..i,|  i.land.  'iliK  c.e.  The  lari.'e  lilciid  siipjilies  ol'  the  tliymid  irlaiid  and 
.if  the  kidii.'y  are  the  most  slrikiim  results  nf  ihesc  o'isci  vatioiis. 

P.v  the  Use  of  the  cahirimeter  method  the  lil Illow  thnni'-'h  the  hands 

iind  feet  of  a  liealtliy  yonnu;  man  has  lieeii  found  to  lie  alioiit  1:?  >:rams 
[Ml-  1(1(1  c.e.  of  hand  per  miiiilte  for  the  ri'.dit  hand,  and  alioilt  half  a 
•.;i;iiii  less  for  the  l.d't.  The  footllow  is  only  alioiit  oiietliird  to  onedialf 
!h;il  of  the  hand  per  1(M»  e.e.  of  tissue  a  dilVereiiei'  whicli  is  laru'cly 
..■■  ill','  to  the  irreatcr  proportion  of  skin  and  the  smaller  i)ro]>ortion  of 
h..nc  in  the  h.ind.  Tlie  averaiie  footllow  or  handllow  for  a  u'iven  imli- 
■•  i.huil  under  ordinary  conditions  is  reiiiiirkalily  constant  from  time  to  time. 
'It   it   is  extraordinarily  seiisilive  to  cliaii'jes  in   llie   temperature  of  the 

■  ■  '.  iioiiiiiciit  in  which  the  siiliject  has  hecn  livinu:  for  some  lime  previous 
!  I  the  iiicasiiremeiil.  In  one  indix  idiial.  ulieii  the  room  teiniicrature  was 
-"  ('.,  the  (low  in  the  riiilit  hand.  eNprcs.sed  in  irraiiis  of  lilnod  per  HMI 
rr   of  hand  or  foot,  was  10,1:  when  it  was  22.8     C,  the  tlow  was  12. S; 

■  1  .11  it  was  2.")     ('.,  12.1;  wlieii  it   was  :'.(>    ('..  IS.").     On  ac uiit  of  the 

hiciu f  temperature  on  the  How.  it   is  extremely  important  that   the 

!.<..!Miieiiients  should  he  made  in  a  small  room  the  temperature  of  which 

ki  |it  ciiiistalit.  or  if  it  must  he  inade  in  the  w  arils,  the  lied  should  lie  sur- 

'  iii'l.il  li\-  curtains.    The  iiieasiiremeiits  made  on  the  hands  of  dispi'iisary 

.    tiihts  shortly  after  coiniiii.'  in    from  outside  air  are  very   likely   to  lie 

'n.Miiiiis.     The  importance  of  makiiic;  such  liloodllow    measureineiits  in 

'•  .liiiic  will  lie  Jilluded  to  later. 

n  c.nirse  the  measurements  made  hy  the  ahnve  method  in  man  tell  us 

■■■■■  v.itr  of  flow  in  tlio  perii>!iery  of  the  1.n<ly.  an.!  furnish  us  with  no  in- 

inn  reyrardinir  the  How  of  liloo.l  throiiLdi  the  \  iscera.     It   is,  how- 

■'.  uell-estalilished  fact  that  the  liloodllow  in  the  central  i)art  of  the 


i 


•-'I-.' 


I'lli:   rlKii  I  AlldV   UK    iin;   iiI.ihih 


I'iiciiiiiliiiii    is    Miiiii'   (II-    Ion    ii'i-ipi ill    with    ||i,it    .'it    tiic    lUM'iplicrv.    .•in 

ilicrt'.'isc    ill    the    (iiic    plncc    lifiiiu'   ;ii'i'(iiii|).iiiii'(|    liy    a    ciii'i'fspiiiiiliii};   <|i' 

Cl'l-JISC    ill    till'   otlici'. 

The  Visceral  Bloodflow  In  Man 

Tlic  viscciiil  lijcHMJIIdW  ill  iiiiiii  (•Mil  lie  iiiciisiiicij  iiiiliicctly  in  tlio  ciiv 
of  the  lllMl.'N.  cither.  (1  I  iiy  liliili'i;:  tlic  i|llillll  itv  (if  oxv'TCil  illpMirlicd  Itv  til. 
IiIimhI  (liiriiic  nil  iiilciviil  >>\'  lime  tli.it  is  less  tliaii  that  fciiuifcd  for  tin 
iiiiKid  til  travel  iiiiee  1(11111(1  the  eirciilatidii  ((id  secdiiils)  and  e(iiiipariiiL' 
this  «  ith  the  (i\\  '.'(•11  ciiiiteiit  of  saiii|iles  (if  arterial  and  xcikiiis  IiIikkI,  or  i  •_' 
liv  canviiii;  a  persdii  td  hrealhe  a  kiid«ii  i|iiaiitity  i>(  nit  nius-oxide  ^as  aiiil 
then  i  idiii^  the  (•(inceiitral i"ii  dl'  this  i:as  in  the  lihiod  al'ter  leaviiiir  th. 
Iuii«:s.      In   the   t'drnier  iiiethud   tlie  dill'erenee   in  iixyjieii   ))erccnlas:e  li.- 

t«('('ii  arterial   and   veiidiis  lil 1   will   he   less  for  a    niveii   alisorptidn   i.i 

iixyi;eii  fidin  tlu'  alveuli  the  iiidi-e  rai)id  the  eireiilation  of  Itldod  tiirnnv! 
the  liinu's.  and  in  tlie  latter  niethdil  fur  tlie  ahsorptidii  of  a  <riveii  aiiidnii' 
df   nitrdlls   (iside,    the    less   will    he   the   Cdiicelit liltidii    (if   tliis    u'as    in    tli. 

Iilddd  the  iiKire  rajiid  tl iiciihitidii.     Olividiisly  these  estiiiiatidlis  inns' 

lie  made  only  over  periods  of  time,  less  tliaii  tliat  taken  for  any  o! 
the  lilood   to  ediii|)lete  one  eirenit    of  the   eirelllat ion. 

The  methods  are  admittedly  oiil.v    aiiprdximate,  lint  the  results  are  '■< 
inueli   interest,  iiiMinly  lieeaiise  of  the  iiidieatioii   they  irivo  lis  ils  to  lli. 

amount   of  hi I  pmii))ed  out   li.\    the   venlriele  with   eaeh  lieartheat.  •>' 

.Inriii!.'  a  ".'iveii  jieriod  of  time.  The  results  liavo  lieen  found  to  v:n 
ednsideral)l.\  ;  thus,  one  ailthdr  iKrdjrhi  places  the  output  of  hlood  )h  . 
minute  as  hotwcen  •_>.><  and  S.7  liters,  wliicli  wiiuld  correspond,  al  • 
(itilse  rate  of  70,  to  an  output  per  lieartlireat  of  from  40  to  120  ('.e.  .\ 
immediate  ami  ver.\  marked  increase  has  lieeii  found  to  occur  durin. 
muscular  Wdrk.  r..\  cdiniiarinu'  the  hhiodlldw  throutrh  the  liaml  \\i'i 
tli.it  tlirdii-rh  the  luiiiis.  ;iii  estimate  can  lie  formed  in  a  ifiven  individu;: 
as  to  the  rclati\c  mau'iiitude  of  the  |ieriplieral   and   visceral   moieties  ■ 

lilodd.      liitei-estiiii.'   results,   wliicli    will    lie   referred    to   later,   lia\<'   li 

dlitaineil   from   such   measurements. 


The  Work  of  the  Heait 

^lejinwhile  it  is  of  inteicst  to  imte  that  we  may  cileiilate  from  ti 
ventricular  output  of  the  hlood  Ih,  nmninil  <if  inirk  llml  Ih,  hnirl  Is  ihii 
ill  iihiiiildliiiiifi  till  lir.iiliitiiiii.  (tf  course  the  calculation  is  afiaiii  oi 
api)r()xiniate,  since  we  luive  to  assume  certain  tisrures.  If  we  assume  th 
in  a  70kih><rrani  mar,  the  »|Ur.iitity  of  l.hidd  is  4."J00  c.c.  i.sc,.  p„jr,.  s" 
mid  tliat  it  takes  ahoiit  one  minute  for  all  the  hlood  to  complete  a  <■ 
eulatioli.  then   the   work    performed   liy   the   left    ventricle   in   one  mini. 


rati:  ok  m<)Vi;mi.st  of  tiii:  iii.»miii 


'2V.\ 


ill  lie  <M|iial  to  that   (lone  in   niisiiin  tlif  al>iivf  (|Uaiititv  of  lilotul  to  a 
In  iL'lit  ('oriM'siMiiKliiiir  to  the  iiicaii  prt'ssurc  in  tlic  aorta.     IT  we  takf  tliis 


•ii'xsint'  as 


|:!()  iiiilliiiu'tcrs  tif  iiHTcni'v,  wliicli   would  poii'cspoiKl  to  a 


iiiiiii  ot"  !)lo(«l  l.T'i't  iiictcrs  lii!:li  i  lo.')  x  1:!0  ^17">."i  uini.  h 


<l. 


1, 


liM-i,  tlio  work  (lone  liy  tlic  left   ventricle  would  lie    l,7')ii  x  4.2=7.^7 

.iliiu'riiia-iiieters  in  one  minute,  or  in  twcntv-t'our  hours  I'ou'.'lily  nhont 
loiloo  kildnrain-iiM'ters.  The  work  done  \t\  the  riirht  ventricle  is  )irolial>l\ 
ilidiit  ohe-tliird  that  of  the  left,  this  licintr  about  the  ratio  of  the  pres- 
-iiies  in  the  two  chainlicrs.    The  total  work  ol"  the  two  ventricles  is  there- 

■  .re  aliout  14fMM)  kilotrrani-iiieters.     This  re|)resents  an  enoruHMis  amount 

■  I  vMirk;  indeed  it  lias  lieen  computed  that  it  is  suHicicnl  1o  raise  a  man 
of  "tl  kilie/rams  to  ahout  twice  the  height  i.l'  the  liiirhest  skyscraper  in 
\i w    York.     The  work  thus  expended  in   I'dit-iin:  the  IiIoimI  Ihroiiirh  the 

.liiillaries  hecoines  cimverted  liy  t'rii'tion  in  the  small  Idoud   vessels  into 
iiial.  the  heal   c(pii\alciit   of  the  ahovc  amninit    nf  ucirk   hciiiji    rouirhly 
'■iilt  :!."i(l  calorics   i  see  p^iyc  .">:>"). 

THE  CIRCULATION  TIME 

The  circiihition   lime,  or  the  time  taken  li>'  a  drop  of  lilond  In  travel 

rtuceii  two  piiinis  in  the  circulation,  can  lie  di'termincd  in  lalioratory 
iiiiiiials  hy  a  variety  of  methods,  all  dcpeiidinjr  on  the  iii'inciple  uf  sceiii;; 
I'lW  Innir  it  takes  for  a  drop  of  some  sulistancc  injected  into  an  artery  to 
ii|ipciir  ill  tlie  eorrespondiiifr  vein,  h'or  cxaiiii>le,  to  determine  the  tiiiK' 
'ilvcii  for  a  drop  of  idood  to  pass  from  the  .iii'_'ular  vein  into  the  carotid 

I 'liy  in  a  raliliit,  a  solution  of  mclliylciie  hlue  in  isotonic  saline  is  in- 
.!''ii'i|  into  the  formec  vessel  and  tli oment  of  its  a|)pcarance  throiiirli 

ill'  walls  of  the  artery  deterniiiied  hy  a  stop-watch.  If  the  walls  arc  too 
''  ick  to  ailiiiit  of  the  cmiiloyment   of  this  method,  a   stronj:  solution  of 

•"liuiii  chloride  may  he  sulistitilted  for  the  methylene  Mile,  and  the  mo- 
I  lilt  of  its  appearance  at  another  |)oint  of  the  circulation  deterniined  hy 
■    iv\  iii^    tlr       '     '   '     '  '■    '■-  -' 


I'M  Ol  lis  apjieai aiice  ai  anoiiier  poiiii  oi  ine  ciicinaiiou  uenTiiiineii  o,\ 
■''Nerving'    the    electrical    conductivity    of    the    \cssel.      Since    the    coii- 
■'.'•tlvity  of  a  lilood  vessel  dei>ends  partly  on  the  concentration  of  elei-- 
'l>tis  in  the  hlooil  llinvin^  thiou-^h  it.  the  moment   at  which  the  salt 
in  inn  appears  will   he  indicated  liy  a  chany:e  in  electrical  resistance 
'i    V  Stewart). 
I',    ^iich  methods,  it  has  heen  fouml  that  the  time  for  the  i)ulnioiiary 
^!    ihitioii  is  very  sliort  compared  with  that  of  tiie  systemic  circulation. 
i  'iihliit  it  is  usually  a  little  less  than   four  seconds-,  in  aii  averajrc 
'      '  ^i  '^  of  nhout  12  kilojiraiiis.  it  is  nhout  ei^fht  seconds;  and  in  man 
'    -  '"inpiitod  to  he  ahout  (iftcen  seconds.     On  the  other  hand,  the  cir- 
'  'i'U  time  in  such  vi.scera  as  the  spleen  and  kidney  is  ri'lati\el.\   hm^, 


•JU 


riii:  <  iiii  I  I  ATI) IN  III-'    iiii:  iii.immi 


h 


mill  iiMUr  MiM'i'|ilil>li'  lli.'iii  iIimI  III'  till'  liiiiu's  til  ililVri'i'iit  I'liiiililimis  m 
lriM|irr'iitili'i'.  Ill  ;i  iloLT  ill  wliii'li  tilt'  |iiiliiiiiii.'ii'y  riri-iiliitiiiii  liiiii'  ^^^l^ 
:iliMl|t    S,.')   sifiiliils,   Ihiit    nt'   till'   splci'll    WHS    illinlit     11    mimmhIx,   ilMil    lit'   ill' 

kiiliii'\  iiliniil  17. ."i  M'l-iiiiils,  Till'  sliiiiti'".!  ciii'iilMliiiii  tiriic  iil"  all  is  "i 
ciMll-M'  tliiil  ill  lllr  rill  oliiiiv  iillil  V  ,  lillt  lli.lt  t  liliiimll  the  rclilia  r.'lll  lint  t'all 
r.'ir  lii'liiinl  it. 

'I'll  ih'ti'lliiilii'  //m  liitiil  I  iniihil  iiii)  liiiir.  \\  r  l\n\\\  klliiu  :  ll  I  lie  a\  rfil'j.- 
.'iiiiiiiiiil  111' liliMiil  p.-iNsiii',' li\  I'iicli  |iail  ilia  iriMii  liiiii'.  ami  f'Ji  tin' a\  I'i'a'ji- 
cit'i'iilat  inii  tiiiii'  111 'rai'li  |iMi't.  Kiniii  Midi  i'<iiii|iiitat  iiiii>i,  wliii-ji  Imwi'Mi 
afi' iili\  ii'iislv  Milijri't  til  riiii>iilrralilf  rriiir,  it  lias  liciii  ici'Uiiiii'il  that  tlir 
tiilal  I'iifiilaliiin  liiiir  in  man  iiiii>l  lii'  mhiii'W  liiTf  lii'lwi'rn  1  anil  1  2-'> 
lllinillrs. 


MOVEMENT  OF  BLOOD  IN  VEINS 

Hrl'iiii'  Iraviiii;  llii>  |iail  nl'  mir  siilijii-t,  a  I'l'W  wunls  iiicx  In'  saiil  rmi 
ri'|-niliir  //"  Jiirrtx  i-mni  nn  il  in  Ihi  iimri  nii  ill  nf  hlimil  in  Ihi  niiis  t'l'mii 
till'  i-a|iillarir^  tu  tia'  licart,  I'.y  tin'  liiin'  that  the  vi'iiiilcs  arc  fi-ai'lifii. 
iiuill!.'    til    tlirtiuli    in    tlir    raiiillal-ii'N    the    hliiiiil    will    have    Inst    innst    lit"    111'' 

rmc-i'  iinpaiti'il  tu  it  hv  tlii'  hrait  arlimi.  Ni'\ citlii'li'NS,  tliis  ri'maiiiiiiL' 
I'ls  II  hri/ii  must  1)1'  I'liiisiili'icd  ;is  till'  liasic  cause  t'nr  the  iiiovcnicnt  ni 
the  vciKiiis  hhiiiil  iirar  tlii'  iit'iiplu  ly.  As  thv  vi'iiiili's  tret  larfjcr,  tw" 
ntlnr  l'ai'tiii>  iniiic  iiitu  |'la.\  :  the  iiiassa^iiiir  iiillilcMcc  <il'  the  liillsclr- 
ainl  thr  \;il\is  iif  tin'  \i'ins.  \',\  till'  iiiii\  I'liii'lits  III"  the  lillisch's  tin'  vciii^ 
whii'li  lie  lirlwi'in  will  he  ih\  thniii-aliy  i-iini|iii-.si'il,  and  this  will  tnul  I" 

i-aiisc  the  111 I  to  111'  mm,  I'll   t'lirwaiil  ami   hai-kwanl  in  tliciii.  the  liai-lv 

vvaiil  niii\  riiiint  luinLr  hnwixcr  incv  I'liti'il  liy  the  ujn'ratioil  ul'  the  vahr- 
Whi'ii  till'  I'liiii'ily  111'  till'  miiM-lfN  is  Milnniinial.  as  in  cniKlitiiiiis  iit'  ii 
lli'alth,  till'  alisriii'i'  111'  this  iiiassaL'iinr  ai-tinli  IH'i-mils  the  lilmul  Ik  stav 
natr  in  the  \i'ins  ispci-ially  in  tlinsc  nt'  the  Idwci'  cxticiiiilirs  in  llpl'iv'l' 
animals,  with  tin-  cnnsi'iiiii'ni'r  that  the  M'iiis  licriiiiM"  ililatcil.  parlii'iilai! 
.jilst  aliiiv  (■  till'  \  al\  IS,  thus  i-ausiiii.'  tin'  rninlil  imi  kimw  n  as  \  arii'nsc  vciii  v 
As  till'  thiira\  is  appruarhnl.  twn  utlicr   t'ai'tiirs  liirnmi'  i>pt'i'ali\  i':  tl' 

aspiiatin'_'  inlliu'i t'  tin'  thmax  •Iiuiiiii'  iiispiiatinn.  ami   the  ni'irati> 

iiilra\i'iiti  inilar  ini'sMiri'  i  si'.'  p.i'ji'  l.'rj^.  'I'lit'i'i'  is  nn  ilmilit  that  l! 
t'liiiiH'f  111'  tlii'sf  is  (if  ciinsiili'ialili'  impiirtanri'  in  maintainin<,'  the  \i-iiii< 
irluin  ui'ar  the  heart,  I'm-  allhiiiiuh  the  ehalitre  nl'  piessufe  imliiei'il  liv  i: 
spiratiun  ainuuiits  tu  milv  7t  niiliinirter's  uf  niereury,  \  et  it  aets  - 
sluwly  that   it   piiiiluci's  a  i-niisiileialile   inlluenee.     The  aspiratiu"?  effi' 

nf    tlii-     \r!i!!-ii-!r    ;i1     tli"     !"'ui!:i-in'J    nf    <!iast"!e     is.    hn\\e\rf,    (if    nil    s:. 

nitie; ill  atli-aetiii'^  hi I  tu  the  heart,  foi'  althiiui.'Ii,  as  we  have  sei  i 

it  may  lie  (•(iiisiileralile.  yet   it   Lists  fur  su  slmrt  a  time  that   it  cuuld  U'  ' 


I  i 


KA'I'i:   IlK    M"V1  Ml  NT   OK   ■mi;    lll.iMili  Jl.) 

,,,,n'HMif  till'  iliritiii  "f  llic  fciliiiiin  i>(  III 1  ill  till-  \riin  fiiMi.      Kvcii   it" 

!l.c  iicuiiliM-  pnsMiri'  <li<l   l;i>l    <>'>■  n  l'>iii:cr  iifriuil,  it   rcnilcl   imt   iiltrart 

Mi.ji-.'  lliiiii   a   -1111111    iiiiioiiiit    lit'   111 I.   liriMiiNf   it    \Miiilil   i-aiiM'   tlic   tliiii 

ll«,  111'  llif  \iins  til  riiiiii'  tiiu'illKf  aiul  tliiis  lilm-k  llio  pas- 


,'lpSlllll'    WII 


lu'i'  tiiwaitls  the  licait. 


f 

■ 

t 

1 

1 

. 

1 

i  .i' 


^  . 


H' 


r:i  I; 


rii.\ni:i{  x.w 

Till-:  (  (•MI{o|.  nl'  Till.;  ciun  l,ATI(t\ 

'I'lir  ii\ilil;ililc  l.liMiil  III  I  lie  \„„l\  is  |iaii-<-l('i|  nlit  In  llic  XilliiHls  iPlu'iiii- 
Illhl  ti^^ll(•^  iicciiiilini!  t..  ihi'ir  iiliiliM'  iirlix  ilicv,  .-iMil.  sjiu'c  tllcsr  villv 
iViim  tiiiii'  (..  tiiiii'.  till'  i|iirsli(.ii  iiiJMs  ;is  to  the  n.iliiif  of  111.'  iiufliimiMii 

""•  •" Ililliisiiis  itiMilM'd  ill  LiiiiirillL-  mImhiI   Ihis  iiiljiistinrlil.     Tun  piissilil. 

IiiciIicmIs  III'  iiiciciisiiiir  till'  >il|i|il\  ;iic:  nil  iinrmsi  in  llu  i/ii/.w  iiiuri  im  iil 
III  till  III!  hliiiiil  III  I'ii'i'iilatioii.  mill  II  I'll  ifiiii,  III  mljii'iliiii  III  nf  th,  n sislnnt  • 
III  llii    jliiir   ill   ililfiniil    riisiiiliir  nniix   Iii(iii!;]i1    .'iliinit    liv    Misnilihitatioh 

ill     nIH'    mill     MI-Mli stli('ti>i|l     ill     nllicl-s.        Until     nf    tlll'M'     IllctlllxU     inillll' 

ii|i('ral('  Imjcf Iicr. 

TW"!     aj.'«'lll'i>'^     I'illl     111'     tl|n||..r|||      ,,f     a-.     I'(■^|)(l||'^illll'      Inl-     |pli|l;.'il|ir    allllUl 

till'  aliiiM'  cliaiMjcs:  i  I  i  .'licmii'al  ^llll^1aM^(■s  iii-  hiiiiiniiu's,  pfcscnt  iii 
tilt'  IiIimmI.  ami  I 'J     till'  hi'iMiiis  s\Nti'iii. 

I  III     intliliini     III    iliiiiiiiiil    M(/iv/,;/,f  (  ■.,    ,,/'    lliiliiinii,  ■> ,     I  |iaj.'i'    ""_'!•)     ill     I  !■  • 

rinilriil  III  III,   rirriiliiliiiii   is  uiiclipiilitcili.x    all  iiiipi.t'taiit   line,  aiul  oj'  tliii^ 

kli((\Mi    at    lilt'    pii'Miit    t twii    '.'iiiNp-.    may    he    mciitiolicii:      I  I)    Mil. 

■''•■' ^   Mliii-h    .'illi'i-    the    liMlri.iiii .■niH'c'iitiati f    Ilic    Mood,    ami 

I'J'  Micallcil  pics'Mii  ."Jill  ili'piivMii'  siilistaiK'fs,  pi'ii.liK't'ii  citlu'i'  i)y  (|ii>t 
IcsN  !.'liili.|-.,  siii'li  a>  the  ailii'ii.il.  m-  |i\  tlit'  activity  .if  tisMics.  An  in 
cr.'asf  ill   liy<lin<jciii.,ii   ('..ii.'ciitiati.iii   of  the  lilnotl   imt   .iiily  atTccts  th' 

li.'.iitlM'at      >..'    p.ii:.'    Iii^    .    hut    i-aiivrs   ,i    maik.'il    ililatati f   tin-    lil 

vessel-.,  SI)  tlial  lintli  til.'  eelitial  ami  the  peripheral  ehiili'_'es  \vill  lie  sin-i 

as  til  eiie.iiii'a'.'e   all    iiM'i'.'a-..'i|   H.iu    ,,(   l>| |    tliiiiiii;li   the   aetive    mjrai- 

.11'  viM'iis  Thus.  .Iiiiiii','  miiM'iiiai-  iii'li\it.\  nf  the  lej.'  iimseles  thi'ic  w  v 
lie   a    teli'l.liey    tn   all    ilii'r.aM'    in    the    liMliiitreilinli    enlieeilllatinn    nf    111. 

''' 1  'i-  'I   "linl.'.   i-.'siilliiii.'  ill  a   '.'leater  eanliae  iietivity  ami   a   ;ri-eiil.'^ 

"I'ti'i'^li  "I'  I'l 1  tiii-nilirli  the  anil.i.  ami  at   the  same  time  the  vessels  '■ 

the   aetiii'.'   miis.l.'    will    ha\.'   Iiei-nm.'   especially   tlilatcil   lieeaiise   nf  th 

pr.i.liictinii  |,\   thrai'liv.'  muscles  either  nf  lactic  acid  or  nf  earl ic  aci. 

Till-   aitive    iiiiisch'   alsn    prn.liices   such    sulislances   as    iiiiiilaznie.    wlii.' 
have   a    pnucrfnl    \  asn.lilal  iii'.'   actinii.      .Such    sulislances    are    snmeliiie 
eajle.i  (lepressnr. 

Thnuirii  Ihe  hnrm,,!:,-  ,-u„!!-.>|  ..f  the  eiivulatinti  is  Uiiilmihii  liiy  nf  irir;, 
im|.nrtaiiee,  it  is  pr..li,ilily  mii.-h  less  sn  than  that  e.x.-r.'isetl  thrniiirh  //■ 
n(niiU!t  siistiiH,   ami   heie   a^aiii    the   cnnlrni    is   I'ent.'re.i    partly    in    ili. 

I'lii 


n 


Ill       IDNTKOI.   iiK    llll     iM.ill.MMiV 


•JI7 


I  .  ;irl    mill    jiilllly    ill    llic    |li'li|illi  till    ir>isti I',      ''''i'    Mi'lM'   ci.lllliil    111'   llir 

,,iil    IN  I'lTiTti'd   iIiiihikIi   llu'    mi'.'U>  iiiiil   sx  In-    hl•l■M•^.   iiml    llial 

1  Micisi'il  nil  the  Miiiiil  NfHscN,  llirmiuli  lIu'  >"• nl  \  iimm-iiiisI  i  ictnr  iiml 

:i>iHliliitiir  iK'iAi'^. 
Tlir   iii'li\il>'    III'   IIh"   IK'IX) lltcis    rriilii    \»liir|i    llir   riililifli*   iiimI    \iisii- 

nmliir    iinpillsfs    lire    <IiM'liiil'.'<i|    in    innl  inMnl    liy    iilTiliIlt     illljlllUi'S    CI. Ill 
I'.'   rnun   llic  \illiiilis   ri-i;iiinM  nf  tllf  limlv.      Wlicil   H   irlillnl   lii'i-iiMli"»   llHilT 
r\\\i-,  \\r  lllllvl    sIlpltiiM'  that   Htilllllllltiiill  lit'  till'  si'll>iil>    tilii'l's  liiis  CIIIIsimI 

iilficiit  iiiiiMiJM's  til  III'  tiaiiHiiiittcil  111  llic  caiiliac  ami  \  aMHiintiir  cciilcis, 
i|Miii  «liicli  llicv  act  ill  Mich  a  \\a\  as  to  |iiim1iicc  iiicrcaM'il  heart  ac- 
■i,.ii  anil  a  lncal  ililatatimi  nf  the  IiIimkI  m'sscIs  ij'  the  active  iflaiiil.  vvitli 
I  1  rlia|i^  a  ciiiisliii't  inn  nl'  the  liliiiiii   \  om'U  ii|    the  icsl    nf  the  Imily. 

THE  NERVE  CONTROL  OF  THE  HEARTBEAT 
The  Vagus  Control 

\\  llll  rcLMIil  In  I  he  cnlillnl  i'\elci>eil  tiirnlluli  llic  \  a'^ll>  ml  \e.   wc  liavi 

:     cMi|>    M'cli   that    the  eiiltiii'.'  nl'  the   I  w  n   iier\es   in    the   neck   eailx'S   the 

'•III    til    i|iiicki'ii    anil    the    arterial    lilnnil    pressure    In    rise,    whereas    a 

•  iiiiiilatinii   nf  the   peripheral   eini  of  the   Mer\e  causes   the   heart    to  lic- 

■  Mic  sjiiwcil.  if  iml   slnp|)i'i|  altnu'clher.  ami   the  Mnoil   pressure  111   fall. 

T'li   the  iiinre  ilelailcd  iii\  est  iualinn  nf  the  naliire  i>(  the  \ai:us  eniilinl 

'he   lieail,  it    is   liecessaiy   tn   nliscixc   the   cNpiiscil    heart    itself      all    ex- 

Mill  111    which,    fnr    nli\inUs    reasiilis,    can    he    llinsl    siliipi>     pcrfmlMCil    ill 

■  111  hjoodeil  aiiiiiial.  such  as  the  fin'.'  nr  liirlle.  hut  which  can  alsn 
I'l'ifnl'llied   in   iiiainiiials  prnxided   arlilicial    respiration    is  inaiiitaiiieil 

li'    'ji'iieral  elTccI   of  the  \aL'Us  in  Imlh  i.'rnups  of  animals  is  the  same. 
•H'ih   apparent    dilVcielices    ma,\    exist    nii    aceminl    of   the    re]ali\c    ilii- 

■  !,iiicc  of  the  dilTcrcnl  parts  ,,(  the  heart  in  tl ri'jiiialinii  ami  ]irnpa- 

'  '"11  nf  the  heartlicat. 

The  Cold-Blooded  Heart.     If  tlic  \ai;us  ncivc  mi  the  liiiht  side  in  the 

■  '  the  left  nerve  is  iiiacli\e  in  this  aniiiiali  is  sliiiiulated  with  a 
\  liihle  electric  current,  while  siniullanenus  rccoiils  are  hciiiir  taken 
the  I';;"'':'.,  linns  of  the  aurieh's  ;iiid  vciilriclcs  in  the  inaiiiicr  shown 
;!i'    acciiiii)iaii.\in!j;  liaciiii;  i  l'i'_'.  (i4'.  it    \vill  oflcn  lie   fmiml  that    llicic 

'■akeniiiLr  i'(  the  auricular   heats   wilhiiiil    any   chaii'^c    in    llmsc   nl 

'nliicic       If  the   siien'.:tli   nf  siimiilus   is  sniiiewhal    imrcascil,   ihe 

;!ar   heal,   licsides   hecnniin^    vvcaker,    will    alsn    liceniiie   slower,    hut 

iilic   itic    \  eiil  nelltar.   Jltllioii     '.    ;iiso   slnwcr,    iMa>     liccoliic   ilisl i\\ 

^"      -'1.     At   liist    sijihl    this   rcsiiK    iiiav    he   a    lililc   cmifusin^,    liecausr 
il  llll  seem   tfi  indicate  that    llic   \;i'^'us   nerxe   weakens   ihe  auricular. 


'  H 

t  i 


li 


111 


f': 


•218 


■niK   CIKrn.ATIliN    OF   Till-;    111.(11)1) 


11  ^ 


lint    slicML'lliciis   llu'    vciitiiciiliii-   lii'at.      It    is   clciii',    Imw  ex  cr,    tliat    llir 
stii'iiiitlicniiii,'  ut'  ttic  Vfiitiiculai'  ln'at   is  iiicfcly  iliic  to  tliL-  I'acl   tliat  lit' 

cavitv  lias  hccdiiic  licltt'i'  (ilicil  «  itii  lij 1  iliiriiij;  diastcilc  as  a  icsiilt  ni 

llic  slciuiii<_'  111'  llic  ailii<'lc'.     'I'Ik'sc  rcsiills  imlicatf,  tiii'ii.  that   with  weak 
stiiiiiihitiim  tile  \a"iis  I'xcrts  its  iliiort   iiilliifiu-f  milv  on  the  aiiiiclc.     Ii 


ru'    'J.      >i:n;ill.ini.  IIS  ii.i.iiii;-   ffuii  :inriiU'  ,iMi|  Miitn.li-  nf  liirlli  \  li.ait.      IMuciii   tin-  ii    •■ 
till     ».i;:m.    «.i-    -li'iril  .!.i|.    uilh    ih.'    itlut    Ih.it    Ihi'    iiiinnilar    l>rat    ilitiiiui-),ii|    iii    l..ni'    l"it    ii'l 
Iri-nuiuv.    uliilr    tin-    \ .  iituc  nl.ir    licit^    win-    prai'ti*  .illy    iinafTi  rtcl.       (  i'rnm    HowiU'-*    /'/i'm   ..  / 

thi'  stiiiiiilat  inii  is  sti-niii;  ciKiiijili  iiiitii  aiiiicli's  i'.iid  vi'iit  rides  cease  h 
lieat  allii<_retliei'.  and  il'  tiie  stimulus  is  iiiairitained,  the  iidiil)iticiii  may  u'' 
(111  I'lir  a  very  Iiiii<^  time  (  I''iy:.  <>.")). 

I'sually,  even  thnujjh  the  stimulus  is  maintained  the  lieai't  Iiej^ins  t' 


*  i' 


IJK     '■?.  -I'.tTt't    '.f    v.ic'is    vtitniilaltiin    nil    liiart    nf    tiirllc.      Nntr    tfir    after    fffrrt    nf    aiiKim  nt.i' 

lieat   Mu'aiii    ai'tec  a   time,  at    first   iiiily   oceasiiiiially  hut    uradually   lie", 
rapiilly.     This  is  kimwii  as  i  uiiju  nii »(,  and  it   iiidieates  that  the  enerj 
jient   up  in  the  heart  durinji  the  vairus  iiiiiiiiitiiui   has  at   last  overc'iii 
llic    inhihitiii;;   iiiliiienee    u(    the    nerve,    which    is    meanwhile    heciimii 
I'atiuued.     Ail  III'  these  results  eiiiild  lie  (|uite  satisfactoi-iiy  explained  '  ' 
the  assumptiiin  that  the  aetiim  of  the  vastus  is  eonlinej  to  the  siii':- 


Till:  t  iiNiKoi.  OK  Tin;  riiicii.ATios- 


J19 


liji-li,   it    will   If   ii'iiii'iiilififd,   <|(iiiiiiiatt's   liie   lifiit    in    tiic    rest   of  tin- 
.alt.     Tiit'lc  i>  cv  idoiK'c,  Imwi-vcr,  that   llii'  va'„'ils  alsii  diicctly  allVt-ts 


I  he   1 


liytli 
tl 


III    tlll'SO    lol 


the  vi'iitriflc.     It   may  lie  stati'il  as  a  tjfiii'ial  coiiclusion 
llts  that   Ihr   liilhii  iii'i    of  tla    nnjii^  uikjii   tin    hniithitit   I'.s- 


,hiillil  vniliriil  iiiiitii  lliDsi  ixirts  of  llir  oniaii  in  u'hii-h  tin  rhulhmic  poinr 
IS  iiiosi  lilijlihi  ill  ri  lojii  <l. 

I'.isiih's  atTcctiii^r  the  rate  ami  slri'M^'th  of  tlio  hcarthcat.  the  va^iis  also 


MICISI'S   a    coll 


tnij  (III  the  cipiiiliirtiv  itv  nl"  the  canliai-  iiiiiscit 


Thus,  if 

,1  partial  hhick  is  iiiNtiliititl  in  the  turtle  heart  liy  a|>iilyiii(r  a  elaiiip  he- 
tui.ii  the  auricles  and  v.'iitriclcs.  sliinulalimi  ot'  tin'  vaunis  ent'ecMes  tin- 
;iiiriciilar  heat  ami  iiia\-  also  cause  a  eipiii|)letc  lieart-hliiek  as  shn\\  n  in 
ill.'  traeiiiir  re))roiiiii-e(l  in  Fi'_'.  M.  It  is  iiiipdrtant  to  ixiiiit  nut  here, 
i;M\\eve:-,  that  iiinler  certain  cnudilious  the  vau'iis  may  a|i|iear  to  increase 
v.illicr  than  (h-crease  the  coii<lucti\  ity  of  the  tissue  in  the  aiiriciiloven- 


ilniillli  ■  ! 


m  "mm 


♦  _^__^_ 

I     ■     .         Ti.i.  niB    !■.    --h"»-    Ihal    \aKn<    Mini.l.il. nay    .liniiiii^li    ti  ln-mi<^i.'M    fr.iii.    .luii.  I.-    L. 

.   I,  -       It    sl„,»»    llu    .ll.rl    ,.|'    slimul.ilini!    Itif    lift    vaK"«   ""    I'.irlial    ( -'    ll    M'«.k    l'r...Hr..l    ..ii 

1   .1   lailK    l.v   .i|.|.li.,.li..ci  ..I"  .lanip  at   a  nu  al..v.  iili  i.  nlar  juiuti..",      Stiiiiulali.'ii   at     i    il<|.r.  ".  .1 

.     i.humin     a'l.l    iMak.iu.l    till-    aiiri.nlar    .  ....tra.  li..n«    ll"vir    lia.inul     \«iiIi"M|    >Ii.»ii.i;    Ih.  ir 

III,-    r.Mill    »a-    ail    iil.r.av    iii    tlu-    .Let..'    ..f    t.l-<k    «itll    .•i-vitl.M,    ..(    v.iltl  h  lilar    C.Mitra>  Innis 

•      li.,.iiiKi.      Initial   aari.ular   lalr        .<>   pir  tniinil.-.      (I-lmii   C.arr.y.l 

l;,iular  junction;   I'm-  example,  it   has  heeii  ol(ser\ed  in  the  turtle  heart 

li.l   when  a  claiiii)  is  so  li^;lit   as  to  ludiluce  complete  lihick   -that   is  to 

..  to  render  the  ventricle  inactive  while  the  auricle  is  still  lieatini;  at 

usual   rate     stimulation  of  the  va^ius,  besides  causintr  the  aillicles  to 

!     •.iiic  distinctly  slowcil.  may  at   the  same  time  cause  the   ventricles  to 

:     1 1  to  the  auricular  heats.     This  result   is  prohahly  due  to  the  better 

.  I  iMccs  of  slow  beats  (.'ettin;.'  throu;;h  the  junction  than  those  which  are  so 
;  ii|Ui-iil  as  to  crowil  one  anollier  on  llie  narrow  briili:e  which  the  con- 
'till  tissue  olTers  to  their  jiassairc  (h'ii;.  <'7i. 

\  .  ry  important   wtu'U  uas  edlitributed   in   tliis  lieltl  by   tl.   K.   Mines" 

V   -  tly  before  liis  lamentable  death.     He   found  that  the  local   applica- 

!' "I  of  atro])iiie  to  the  sinus  eliminntes  the  clVci't   of  stimulation  of   the 

!■  t-ncraninl)  vagus  on  the  rate  of  the  heartbeat,  while  the  elTeet  on  the 


I  i 


In 


I  \ 


220 


■llli:   tlHCI  I.ATKIN    UK    Tin;    iSI.OOD 


amiiMiliiM'ntiii'iiljif  .jmicliiiii  iiml  mi  the  Nciitriclc  iciiiiiins.  Afti-r  tip 
iilri'|)ini/;itiiiii.  \;i!.'iis  stiiniihitidii  dclnys  the  lriiiis)iiissii)ii  of  licat  fmin 
aiiiii-lr  to  M'litiicit'  ami  slimtcns  tin-  tiinc  of  oach  licat  in  tlu>  vontriclt 
ll  was  initliiT  t'liiiiid  that  li.v  tlir  local  a|)))lii-ati(iii  of  atropine  variouv 
parts  of  the  ventricle  can  l)c  I'cndcred  iiTcsponsive  to  the  intllieiiee  <il 
the  vaiius  and  the  elTects  of  this  iicfve  oii  the  form  of  the  eardio}fifiiii 

1 iitied  at   will.     These   i-esults  have  an   iiapoftant    lieaiinur  in   the  in 

tcrpretation  of  the  canse  of  the  T-wave  of  the  cl.'ct  io-eardiof;raii! 
which  \vill  he  rcfcncd  to  later.  Mines'  icsults  sho\\  that  the  proli^i 
lilc  explanation  i>  that  llie  T-\\a\e  is  dne  to  the  j.'reatei'  diirati  in  of  tie 
excitatory  state  at  the  hase  than  at  the  apex,  for  li.v  alteriny:  the  r<'lati\' 
ihiration  of  this  >tate  at  hase  ami  apex  li.\-  the  aliove  methods,  lie  eonhi 
canse  the  'j'-waxe  to  appeal'  or  disappear. 

The  direct  excitahility  of  the  heart  muscle  to  ext'inal  stimuli  is  als" 
depressed  durim.'  \asiUs  stimulation.     Tiiis  elVect    is.   however,   iiof    e\  i 


\JMM 


m\mv^W0^^   (Ml  iMiiiiiiiiKiiii 


l'-  '"  li.i'iii;  1..  -li..\\  (tilt  v.u'it^  ^ini.iil.iiion  111  IV  l;ti)|it.)tt  ti  .itivnii^Ki..n  trt'in  aitii^.- 
*''l'"l'~  I'  ^li"«-  ill.  ilK.t  ,|  ukIiI  v.ikhs  MiiMiilalM.ii  "ii  lu-.irtM.Kk  |T"iliui-cl  in  llir  liirll. 
.1  .1.1111]..  Ipjiir  li.i.iin:  i,,..i.l«  MMlri.lc:  l..«ir  Ira.  ink',  aiiii.l.^.  W.-ak  (ara.li/.ili.m  ..I  thi-  i. 
i.ii;.!--  II. nc  l..i!iiiiiMiK  .11  .(  .iiri.l.il  tlir  .l.itr.r  ..f  1,1. ,<k  ..iily  al  f,  kIi.m  a  IimkiIiiii..,!  |.i  i 
i.HM.ii  .iiiiutil.ir  iiiiiria.  ti.'iw  i,i.i*..l  a  ..invli  vi  iilri.  iil.ir  i..iitrai  ti.iii  .Al  H  ^Iroiitfrr  fara.l 
I. Mil  ,.1  Hi,,  -aiiir  II, TM-  |.t...lii,,,l  luarki-.l  ~l.,«iiii;  .,1  ili,  niinili-.  in  <i.ii«i  .|iii  n,  c  ,,f  »lii,li  lli,-  L! 
.h-a|.i.,ai,.l  ail.l  tin  \,i.tn.l,~  ,  ..iilr.i,  li.l  allir  i  ,i.  li  .iiriiMlar  ,.,iilr.uli.,ii.  IM.i.k  i.M|.|„ar<-.l  u' 
ill.     Kit,     a«..iii    I.,.. .1,1,    i.i|.i.l.      hiiiMl    ,11111,  liar    ral.  Ii.    |,,  r    iiiiiiiiti-.      i  I'n.iii    ('.amy  I 

dciil  in  the  ease  of  all  hi-jiits,  liiit  is  seen  in  those  of  certain  li.shes  (e.  i: 

tl 1 1. 

The  Mammalian  Heart.     Tin'  ai-tion  of  the  vasrus  on  the  inaiiiinali^i 
heart  nia.\    he  in\  est i.^.n..,!  cither  h.\    I'xposiii!;  the  heart  and  eoiiiieelii' 
the    auricles    ami    \ciitrieles    with    sjieeiall.v    d<'sii.'ned    recordinsr    levr 
(riiyoc.'ii'dio..'iapli   .  or  if  «c  desire  to  study  the  inlluence  on  the  heart 
a  whole.  li\   lakiny  a  lilood-piessiirc  traciii;.'  from  oni'  of  the  larv'e  arteii. 
Ii.\    means  of  a  spriie.'  iiiamimelei'.     The  results  arc  in  -ieiieral  similiir  ' 
those  olisci\ed   on   the   fio^'  of   turtle   jieart,   the   main   elTeetis  liciiij^  ■' 
veloped  on  the  auricle      ( 'oiisidci  jiMc  iJifTcnnees  are  found  in  f!io  etV, 
on  tiie  heart  as  a  xvliolc  in  dilTcrcnl  .inimals.  particularl.N   with  rejrard 
the   fai'ilit.\'  with   which  cMai.ement   occurs.      In  tlii'  dof;  when  the  va;' 


rili:    CON'liiOI.   OK    Tin     clKill.XTInS- 


221 


i^  coiitiiMiously  stiiiniliilcd.  tlic  liciirt  is  likely  to  rciiuiiii  iiiliiliiti'd  Tor  .-i 
I'lrisi  tiliii',  \\  liiTCiis  ill  till'  r;it  the  iiiliiliit  ion  is  very  i|ilick!\'  liriikcii  into 
l;v  I'svapcniciil.  II'  tiic  Irjiciii}.'  is  liikcii  directly  fnnii  llic  lii-iirt.  it  will 
lieiiut'litly  lie  ()liser\eil  in  tlie  dnir  tli.'it,  wlieii  the  ese.ipciiii'lit  (leeiirs  diir- 
•!!!.'  Viifrns  stiiiiilliltion  it  is  only  the  vi'iitriele  that  is  lieiitiiiu'.  the  iiiirii-le- 
-lill  reiiiiiiniiij.'  inliiliited. 

It'  flie  stiiiiiiliitidii  dl'  the  vjilhis  is  discdiitiiuied  after  sniiie  time  in  an 
iiiiiiial  whiise  lilodd  ])ressiire  is  lieini.'  reediiled,  llie  )>iessiii'e  will  Hot 
Hilly  (|iii<'kly  reeiiver.  hut  will  usually  nveishimt  the  imiiiial  level,  mainly 
i.irailse  111'  tlie  asphyxia  wliieli  lias  heeii  produced  dining  the  iieriod  of 
iiiliihition.     The   asphyxia   raises   the  hyilnijieli-ion   eonceiitration    of  the 

M 1  and  this  stiiiiulates  liotli  tlie  \  asoconstiictor  center  and  the  heart 

irtion   I  pa;:e  KiS).     Thi    increased  heart  action  is  also  partl.\   owiiiir  to  the 
liict  that  diiriiifj:  \au'Us  iiiliihition  the  hcatiiiir  pouer  of  the  heart  lieeonies 
ii|iro\cd    I  pajre   '2'2'>) . 

As  an  ollfeonie  of  recent  work,"  it  has  lieeii  shown  that  the  riirht  vau'iis 

i!.!\e  acts  iiiainlx'  on   the  sinoaurieiilar  node,  iilid   the  left   xaifils  on   the 

!(  II  ieulovciitriciilar  luinillc.     This  is  in  ayreenieiit   with   the  oliservatioiis 

i|.  Nriiheil  above  on  the  cold -liloo<  led  heart    (  paue  "JIT".      Stillllllat  ioli  of  the 

■lit  vanns  always  causes  slowinjj  and  wcakeiiiiii;  of  lioth  the  auricular 

iiid  the  ventricular  heats,  hut  stimulation  of  the  left  vat'us  is  sonietiuies 

liivcrxed  to  liave  hut   little  inllueiiee  on  the  auricular  heat,  altliouirli  it 

!ii.i\    pniducc  a   condition   of  partial   heart  lilock :  or.   if  a  clamp   is  ap- 

l.lii'd  to  the  ;iuriciilovi'iitricular  liundle  so  jis  to  produce  a  partial  heart 

\■'^^<■k,  then  diiriii'r  stiiiiiilatioii  of  the  left   va'j:us.  the  Mock  may  lieconie 

'iiplete.     Tlicrr    is.   liowever,   a    coiisideialile   o\ crlaiipiliir   of   thest    in- 

.   lues,  at   least   in  the  ciise  of  the  left    xajrus,   for  this  nerve  also  acts 

iiii  >idcralily  on  the  \eiitricle.  iiillueneini;  perhaiis  not  so  much  the  rate 

:!-  'iic  force  of  the  contrai'tioii.     It   has  1 n   I'ound  e\periiiieiitally  that. 

li  nrder  to  demonstrate  the  sju'cilic  action  of  the  left   vaiius  on  the  liuii- 

'l''-,  it   is  most   suitalile  to  stildx'  the  rel;itioiishi]i  lietween   auricular  and 

'    iiiiiciilar   lieats    when    the    auriide    is    lie,;tini:    rajiidly    as    duriii'j    the 

.  I'plicatior,  of  artilicial    i  electrical  i    sliiiinli   to   it.      (•rdinarily   the   i-on- 

":    lion  produced  li.\-  each  stimulus  passes  into  the  ventricle,  liut  diirint; 

'  iiiiilation  of  tlie  left  vaiius  it  is  I'ound  that  ever\   contrai'lion  does  not 

'    -^      These  ex])erinieiit-.  raise  the  (|Ue>tion   as  to  what   the  iiitluenee   of 

til    nerve  iiia.v  he  in   Mockiiiu:  impulses   from   the  auricles  to  the   \eii 

ies  when  auri<Milar  tilirillat  ion   is  present.     It   miirlit   lie  ex)>ected  that 

■i  ■   '••ft  vaj^us  woiihl  prove  more  elTcctiial  in  this  reaard,  luit  ai'tuall>    it 

'        '"■eu  found  that  liotli  vairi  have  th.e  siiiue  eiTcc-t. 

Action.-   I, ..pulses  are  coiistantl.\'  iiassiiiji  aloni;  the  \aui 


"onic  Vagus 

lie  heart.     On  aecouiit   of  this  so-called   tonic   action,   the  lieait    rati 


TMK   flldl  I.ATION    ^t^•   TIM;    lU.noh 


iiii'l'('iis('><   wlicii    tin iiliiiiiity   i>r  tlic    XiifjlIM   iMTVc    is   IiidUcii   citliff 


iltiiitr  or  liv  frcc/iiii 


liiiiliciii  (if  iit'ivc 


Kiir.  -*li'.     The  crrcct  is  us 


iiifiiiis|iiciiiius  wlicii  iiiir  MiMvc  iiiilv  is  cut,  liut  ill  iiiosl  iii;iimii;iis  it  in 
••ciliics  (|uiti'  in.'irkcil  wlii'ii  lu>tii  iiic  I'lil.  ( 'ii.Mlivrc  in  tiu"  licnit  rale  \ui> 
(liii'i'il  liy  imisfular  clVorl  is  nnii'li  iiiDfi'  jrrii(lii:il  in  aiiiiiiiils  witii  inarki'l 
vatriis  tone,  sncii  as  liiintinyr  ihi<ss.  than  in  tlinx-  with  little  vaj,'iis  tunc,  as  in 
(Idliitstic  rahliits.  T  .c  clcu'icc  nt'  xa^'iis  tone  Ihficl'ori'  hears  a  relation 
sliij)  III  tlie  slavinu'  power  of  the  animal  tor  |>roloni;ci!  iiiiisi'uiar  etVoii 
If  is  usiiall.v  ill  ilevel(i|ie(l  in  eoUl-hloiuieil  aniiiials.  It  is  (|iiile  iiiarUe.; 
in  tlie  ease  of  man.  as  is  e\  iileiit  on  oliseivini;  the  heartlieat  het'ore  aii'i 
after  ui^iiii;  a  suflieieiit  ilose  nf  atropine  to  paraly/e  tiie  termination  oi 
the  vajrns  in  the  heart. 

The  exai't  hieation  of  Ihe  nerve  (ells  tliat    form  the  center  of  (liseliari; 
iiiir   impulses  alony:  the  vayiis  <iiiiTs  to   liie   heart    can   not    he  mach'   oii' 
^^i|]l  certainty.  Ii;i;    they  are  no  doiilil    part  of  tlie  i^rcat    motor  liiich'n-. 
I  aiiihiiruiis"!.    froiii    aImcIi    arise   the   (ihers   not    only   of  the   va^'iis   hut    o! 
till'    ^lossoph.iryii'ri'al    nerve.      The    tone   of   this   vaLrus   center    is   almo^' 
without  ilouht  ile|>eiiilent  upon  the  cou>taiit  transmission  to  it  alouir  tin 
sensory  or  alTcrcnt   liliets  of  impulses  eominir  from  various  ])()itious  <'■ 
the  hody.     .\ccoriliny;  to  the  slrenu'th  or  numher  of  these  iiiii)ulses.  th' 
tone  may  he  increaseil  or  iliminisheil,  thus  alleriiif;  the  rate  of  the  hcait 
It    is  possilile  of  course  that  the  tone  can   Ire  maiiitaincil,  iii(lci)enilenll; 
of  the  alTeient    impulses,  hy  the  ;iction  oil   the  center  of  clioiuical  liiet;i 
liolic  juoilucts  or  hormones  luodueeil  in  the  cells  or  carried  to  them  in 
the  hlooil.     We  know    ;it   least   that,  like  the  I'cspii  atory  center,  that   o! 
the   vauiis   is  exeitalile   liy   such   hormones   as  the   hydroiren-ion   coneei 
(ration  of  the  lilood.    The  tonicity  of  the  \at;iis  center  is.  liowever,  mainly 
dependent   Upon  the  piissaiTc  to  it  of  alTeient   impulses,  and  as  evideiH' 
for  this  coiiclu'-ion  iiijiy  he  cited  the  oliscrvalioii  that,  after  section   '• 
most  of  the  alTeient  nerves  to  the  mcdull;i  (as  hy  eiittilii,'  the  sjiinal  cm  ' 
hiu'li  up  in  the  cervical  reL'ioni,  stihseipient  section  of  the  two  vairi  do. 
not  i>roduec  anyth  ntr  like  the  usual  dci,'ree  of  ehantre  in  the  heart  rai' 

The  Afferent  Vagus  Impulses.  Tlie  alTercnt  vairus  impulses  may  con 
from  ]>ractically  iiiiy  jiart  of  ihe  hodv,  li;i\iii}r  heeii  first  discovered  1 
flic  sinii)le  cNperiment  of  tajipinir  flii'  alulomen  of  the  froi.'  with  the  li.r 
die  of  a  scalpel,  ^\  hen  slow  iiur  of  the  lieart  rate  is  ohserved,  Cuttiiii.'  tl 
v.iL'i  aholishes  tiie  rellex.  Similjir  cardiac  inhihitiou  is  iiroduced  hy  in 
chanii'al  stimulafii  n  of  the  tail  or  !.'ills  of  ;in  eel.  \t\  iiuimiiials  stimiil 
tioii  of  the  cenlijil  end  of  any  sensory  iiervC  usually  slows  the  Ina 
liioiii.'h  somel  lines  Ihe  iipposjfc  cfTecl  occurs.  The  |)uImouarv  lir;ilich 
(if  ihc  \ai.'ils  ;ire  particularly  sensitive  in  )>roduciii<;  rellex  inhihiti" 
and  distinct  results  are  iisiiiillv  olitained;  hv  stimulation  of  the  tcrinii 


m 


Till-:  coNTKoi.  (IP  Tin;  ciRcri.ATioN' 


111. MS   111 

:i<  liv  in 


tlic  (iftli  nerve  in  tlie  inuedsii  of  llie  iippe-  lespinitorv  piissaues. 


Inilii 


ijf  iiiniiiiinia  vaixir 


1) 


stiiiin 


liiti 


.(•  tl 


le  sciisorv  nei've  cm 


!j:s  in  the 


].l. 


iiii'vnx,  as  liv  swailowin 


L';  ami  of  tlie  inueosa  of  Ilie  larynx. 


when    a   sn 


l)Sli 


'swalloweil   tlie   wronji   way 


Tlie    sensofv 


)f  tlie  alxloiiiinal   viseera  seem  to  lie  ]iaftienlaily  aetive  on  IIk 


MLMis  eeiiter.  as  is  seen   in    ii'ritation  of  tlie  sei 


)f  til 


stom- 


su<'li  as  occurs  ill  u'nstiitis. 


I'rofouml  iiiliitiition  m 


ilso  I 


le  caused 


ileiit  stiiimlation  of  tlie  mesenteiy.  as  from   a  lilow  nil   the  alitlo- 


II,  or  l.\'  irritation  ol 


tl 


le  sensorv  m 


rves  of  tlie   intestine,  either  tm 


•liiinical  or  lieeaiise  of  disease.     AiKither  iiiteiestin<,'  ilhistrMtioii  of  affer- 
ht   v„i.'iis  stimulation  is  olitained  hy  inessiire  on  the  outer  canthus  of 


till'  eve. 


Ill  siiiiie  iiK 


This  ocnloiiiotor  vasrns  reflex,  as  it   is  called,  is  very  mat 
lividiials. 


iked 


Tliroiiu'h  which  of  these  afferent  paths  it  may  he  that  the  constant 
iiiiiili  are  transmitted  to  the  vatrus  center  to  eiialile  it  to  maiinain  its 
:ic,  can  not  lie  said,  although  it  is  very  likely  to  he  thrmi'ih  the  vis- 
■rii!  nerves. 

Ill  coiisideriiiK  the  cause  for  an   ohserved  chau<ro  iu  heart   rate,  wc 

1  the  noss'.iilitv  that   th"  action   may  have 


^t    of 


course  hear  in  iiiiiu 


d,  not   throu-ih   the  vasriis  center,   hut   throiijii.   the  sympathetic 


ilrr.     Thus,  when  the  heart  1 on 

diiniiiution   in   the   vajjus   tone  o 


's  (piicker.  it  may  he  owinir  either 
to  an   increase   in   the   discharffos 

h 


.'_'  the  symi)atlietic  nerve  from  the  aiiirmentor  center.  That  .such 
i\  action  throutrh  the  au^mentor  center  does  occur  under  oxperi- 
ital  coiiditions  has  heeii  clearly  shown:  for  oxami)le,  if  both  vasrus 
'.  :ire  cut  and  the  peripheral  end  of  oiio  of  them  stimulated  niod- 
•  1   .  -'.  as  to  hohl  the  heart  at  about  its  normal  rate,  the  stimulation 


.f 


cciliiiii  sensory  nerves  may  cause 


•rease  in  the  heart  rale.     Reflex 


itlictie  control   of  the   heartbeat    is   however   no   doubt   much   less 
ipiiit.iiit  than  control  throujrh  the  vafjiis  center.     When  it  docs  exist 


ii    III 


that   the  actual  rate  of  the  heartbeat   at   any   tiiveii   moment 


-I   r 


•piesent  the  al^tebraie  sum  of  two  op|)osiii};  iiifluei 


with  Miat 


iiriis  preponderatiiisr.     The  advaiitatre  of  such  a  double  'i'- 

tliat   it   insures  lu-omiiter  adjustment  of  the  boat.     If,  foi   ex- 


tor  any  revsoii  ipi 


ickeninjr  of  the  heart   rate  is  necessary,   it  is 


'■■  'I'.'lit  about  most  i)romptly  if  the  vairiis  tone  is  dimiiiished  at  the  same 
1'  iiii'iit  that  the  sympathetic  ttiiie  is  increased.  Such  reciprocal  action 
'  iiitiiu'oiiistie  inlluencos  is  the  usual  rule  in  the  animal  oeonomy.  Thus, 
v'l'ii  the  knee  joint  flexes,  it  does  so  not  merely  because  stimulatiiii; 
ii'i>iiises  arc  transmitted  to  the  hamstring;  muscles,  but  also  because  at 
ti''  same  moment  inhibiting  imiuilses  are  transmitted  to  the  extensor 
^■li's  (see  jiafie  814). 


!     h 


'A 


Tlir    I  llfil  l,\TMi\    n|-     llir    lll.onh 


Scxciiil  |)(issiliililics  li;i\i'  In  lie  ki'pt  ill  iiiiiicl  \\  III'!!  >\c  iiH('iii|)t  t 
ili'tcriiiiiic  III)  i.iiilnnf  niiisi  luruK  niisi  iri  il  ilaiiiiii  in  llii  luorl  nili 
iii'iii.  Tlius.  ■,\  >lii\\  iiitr  111'  llic  r.ilc  nuiy  lie  iliic  to  iiii'i'luiiiiciil  >tiiiiiil;il  ini 
111'  tlic  \;iuils  liuiik,  ;is  ill  pifsMilc  nil  the  iici'M's  liy  ii  tlllilni'  of  iinciirivi 
ill  tlic  iicrk.  'I'li.it  siicli  iiiii'liiiiiiiMl  iiiiljiliiiii  iiiiiy  sliiiiiil.itc  Ilic  vjiiiii 
i>  ciisily  (iciiiuiistriitcd  in  iiiiiiiy  iiMli\iiliiiils  liy  jiiiplyiii'j  |)i('ssiir('  in  tl^ 
Miiriis  wlicrc  it  lies  in  the  ncciv  in  fimil  ol'  llic  sivtli  cciviciil  xci'trlpi, 
Siii'li  pifvsiiri'  sciiiictinifs  picpilnrcs  sn  piordiiiiil  ,111  iiiliiliiti<iii  ot'  the  Inai 
llnit    Iciiipnr.-ii  \    l(is>  dl'  ciiiiscidiisncss  dcciirs.      It    i>  iprtcii   iiii   iiiisfit'i'  i\ 

iK'lillK'Ilt    111   pcit'iiiiii. 

•  'liJiiiL'i'  in  till'  lii'.irt   lilt.'  in  niaii  iii.i>    In'  caiisi'il  liy  iliici-t  stiiiiiilatii' 
111'  till'  Viiirils  ft'iiti'i'.  as  liy  till'  picsi'iii'c  ul"  a  tuiiinr  iir  ji  lilniid  clnt   in  ll' 
nii'ililil.i,  111'  liy  till'  artiiiii  mi  tin'  rciitrr  ul'  muiic  iiiiiisiial  liiii'iiiiiiic  in  1I 
lijiiml.      A    L'liii'i  ill    ini'iiasi'    in    iiitrai-taiiiaj    pii'ssiiri'  also  stiiiHilatcs   tl' 
\atnis  I'cnti  r.     Tin'  slnuinu:  nl'  tin'  lii'ait  wliirli  nri'iirs  in  asphyxia  iiiii:! 
lie  lint'  I'itlnr  tn   tin'  ai'timi   nl'  linrtiiunt's   ( liyilrnu'rii-iiin   cmifciitratioi: 
ill   till'  IiIihmI   as   till'   ii'siili    111'  till'  asphyxia,   or   to   thi'   iinTonscd    iiiti.i 
ci'aiiial  |)ii'ssiiri'.     That   the  lattrr  is  tin'  iiniro  iiiipnrtaiit  catlsc  is  slni\\ 
li>   till'  t'ai-t  that,  it'  tlii'  cisi'  in  liitiuil  prtssiiri'  is  pii'vcnti'il  liy  ciiniicrt  in 
an   at'tciy  \\itli   a    iiii'i'i'iiry   \al\i'.     that    is,   with   .-i   tiilic  (lip])in^   iiitn 
i-yliiiilcr    111'    iiii'iciuy    to    a    ih'ptli    cm  ri'spninliiisi-    tn    thi'    iioi'iiial    lih"' 
pi'i'ssiiic,  sn   that    when    tlif    prcssiiii-   tcmls   tn   risi'   the   lilnnd   csi-aiii's, 
llii'  sjiiH  in^'  111'  till'  hf.-iit  is  iini  nliM'i-M'il.     Till'  cxrit aliil ity  nt'  the  aft'i'ii': 
vairils   liiiiis   ill    tin'   liliiL's   is   mi'atly    iiii'ii'asi'il  iliirim;   thr   railii'i-  sla- 
nt' rhlnrnrmiii  ailniinistratinii. 

I'inall.v    it  sliniilil  111'  pninti'il  mit  that,  allhniiyh   w  r  have  iin  vnliiii1:i 
rniilinl    nl'   till'    artixily    nl'   tin-    vatriis    I'i'iiti'i',    its    ai'tivitics    arc    siiliji' 
tn  u'icat  \,iii;itinii  as  a  ri'siill  i\\'  impiiisi's  tr,'iiisiiiitti'il  t'loiii  ct'iitci's  hiirli- 
np  in  liii'  riTilii'iiNpiniiJ  a\is.      It  is  hy  llii'  npiTatinn  mi  thi'  vajrus  ri'hti'r 
siirh    iiiipiilsi's   that    rlian'_'i's    in    hi'ait    latr   iii'ciir   (liiriiitj   t'lnntiniial    '■ 
i-iti'ii    lit,   t'liuht.  iti'      Thr  iiiiTi'iisiil   hi'iiit    rati'  in   iimsciilar  I'xi'i'risr 
pi-nlialilx   liiprinli'iit  iipmi  a  iiiiiiiliir  nl'  i-aiisi's,  siii-h  as  the  iri'mliatinii 
till'   liiiitnr   iinpulsi's   mi   In   thr  iMi-ijiai-   ri'iiti'ls    I  SIT    p,ii;i'  412).   tin'   li-r 
ti'iiipt'i;itnii'  anil  I'haiiui's  in  thi'  hyilintrcii  inn  I'mn'iiit latinii  nl'  the  liln^ 

Ctl'. 

Mechanism  of  Action  of  Vagus  on  the  Heart.  I'liysinlnulsts  havr  i  . 
nraliy  lirrii  riiiii'Us  as  In  thi'  i'\ai-t  iiianiii'r  in  whirli  the  vairils  ini 
liiiiiL's  ,ilimit  iiihiliitiiiii  nl'  lirart  ai-tinii.  Siiiiilai'  iiihiliit  imi  as  a  ri"-' 
nf  stiiiiiilatimi  nf  i'lTi'r"nl  nerves  exists  in  the  ejise  nf  the  ililatm-  (ih. 
tn  the  lihmil  vessels  i  pa'_'e  "Jill  1  ailil  the  sy  lllpat  lilt  ie  llel'M'  tn  the  il' 
tine  I  pajre  -1(17  l.  Inhihitinll  ni'  Vnlllllt.liy  inllyeles  e,lli  he  plnillleeil  .^ 
tliiniiL'h    the  eeiitral    nervniis   svsteiii    Ii\    st iiiiiilat inn    nt'   alVeretit    mi 


,Kir,ht  .hil  r. 


ProudiKfhonic  vvuren 

i'oitq^iiiifliunic  neuron 


iiif  Vi'iii  cav'j 


HouM  /7om 
Ht\irt/vtcr 


Bidder's    (janqhtjn 
n  Mr.Qulo  ventricular  junction 


in  imojuricuiar  junition  [Crt-ni'iit] 

thetic  fibrvi-  do f ted  lines 


,11      '. 


I  ( 


I  * 


Tin:  coNTUoi,  OP  Tin:  circii.athin 


( |iii(rc  S14).  It  is  nut  the  ihtm'  lilicis  llicinsrivcs  tluit  :in'  rcs|)(iiisilili' 
for  t)u!  inliiliitoiy  clTi'cl,  ftir  it  lins  liccii  t'liinitl  tliiit  if  tlic  ixM-iplici'iil 
rinl  i(f  n  rut  viisiiis  iKTM'  is  (•(iiiMccli'd  witli  till'  contrjil  fiid  of  "lu'  nf 
till'  iiiitciior  I'ddts  (if  tlic  cfrvicfil  |)iprti(in  of  tlio  s|)iniil  rortl,  llio  axons 
Ml  tiie  liiltcr  wlicn  tlicy  <;io\v  down  into  the  vastus  tiimk  during  tlio 
ii'ircncratioii  wliidi  follows,  stinnilation  of  tin'  i't'!.'<'iH'rat('d  fillers  will 
still  priwlucc  inliiliitiiin  of  the  licart.  Tin-  nature  of  tlio  fillers  ean  not 
therefore  lie  the  faetor  upon  wliieli  the  inhiliitinj;  influence  of  flie  vai;tis 
IS  dependent.  This  leav»'s  the  terininjil  apparatus  of  the  vasius  fibers  in 
ihe  heart  as  the  struetures  in  wliieli  the  stinndus  eonveyed  fo  tlieni  is 
iindered  inliiliitory  in  nature. 

Tliero  lins  been  eonsideralile  spoeulalion  as  to  -what   kind  of  elianfro 
iinist  bo  oeeurrinjt  in  tlio  heart   in   order  to  eause  the   inhibition,  but 
I'lactienlly  nothins;  that  is  definite  is  known.     One  sijiidlieant  faet,  liow- 
i\iT,   is  that   the  eleetrieal   <'Urrent   led  ofl'   throuith   nonpolari/.able   elee- 
tiiides  from  two  portions  of  the  auricle  one  of  which  is  injured,  does  not 
i.ike  the  same  direction  when  the  vajfus  nerve  is  stimulated  as  that  which 
it  takes  when  the  ni<itor  nei've  of  a  similarly  observed  muscle  is  stimu- 
lated.    A  positive  instead  of  a  negative  variation   is  observetl.     Now. 
-iiH'c  a  iu»j;ative   variation   is  always  aecom])anied  by   active   chemical 
Imakdown  changes   oeeui-ring  in   the  muscle   to  supply   its  ener<ry   of 
loiitrnetlon,  it  is  nsstimed  that  the  positive  variation  neeompanyin<f  stim- 
ilation  of  the  va<rux  must  indicate  that,  instead  of  a  katabolic   priM-css, 
a   liiiildintf  up,  or  anabolic  process,  is  bciufj  excited.     This  conclusion 
'  ■uld  fit  in  perfectly  with  the  well-known  faet  that,  after  the  heait  has 
!■■       held  in  standstill  for  some  time  by  vastus  stimulation,  the  beats  arc 
-,.     n,  r  after  the  inhibition  has  passed  oft  than  they  were  before.     The 
\a>;iis  stems  to  have  a  conserving  influence  on  the  heart.     During  the 
iirliiliition  produced  by  it  enertry  material  is  apparently  stored  up  in  the 
litait,  so  that  when  the  beat  is  reestablished  it  is  stronger  than  before. 

The  Manner  of  Termination  of  the  Vagfus  Fibers  in  the  Heart.— This 

subject  is  of  considei'able  i)liarmaeo!o<;ic  and  therefore  therapeutic  in- 
t'  iN'st.  In  approachinsr  the  i)rolilem  it  must  be  remendiered  that  the 
\i!;us  fibers  belon<r  to  the  so-called  cerebral  autonomic  system  of  nerves 
s.'c  page  SS2).  They  ai'c  therefore  fibi-i's  which  have  cell  stations  situ- 
ated near  their  peripheral  tci'mination — cell  stations,  that  is  to  say,  in 
Aliich  ganglionic  niedidlateil  fibers,  by  forming  synapses  arouinl  nerve 
■ills,  become  connected  with  postganglionic  nonmedullated  fibers.  The 
txistcnce  of  ganglia  in  the  heart,  i)articnlai'ly  of  the  frog,  has  been 
known  for  a  long  time.  These  ganglia  are  located  at  the  siiniauricular 
junction,  at  the   interauricular  septum,  and   in   the   ventricle   near   the 


■_'•-'»•. 


I'lll     I  IKc  I  I   v'l'liiN    iiK   'I'lll     hi  iHili 


aiiriciiliiv ciitiirulni  .jimcl'mii.     'I'lif  riiiii-liiiii  ol'  tlu'  u'lHiKlia  is  to  serve  n 
eell  >lafi<ili>  iiri  llie  ciuii-.'  ol'  llie  vau'lls  iiei'ves.      (  Kiif.  ''^.^ 

Nieiitiiie  is  a  111111;  w  liidi  in  eeitaiii  eniieeiitrntiuns,  if  ii|>|>lie(|  loeall.\ 
III  s>  iiipatlietii-  uiiiiflia.  s|iecilii-all\  paralv/es  tlie  svnajises  lietweeii  tli. 
elitis  III'  l)ie  IHeL'aliifliiiiiie  liliers  aiiil  tlie  cells  I'lniii  wjiieli  tlie  pusl 
jralitrlioiiie  lilieis  aiise.  II'  tliis  driii.'  is  applieil  in  a  1  pei'  cent  soiiiliiMi 
til  the  lieai't,  stiiiiiil.iti<iii  of  the  \auMls  tliiiik  no  loriu'er  proi'.uce.s  iiihihi 
tiiiri,  liiit  if  the  stiiiiiiliis  is  ap)>lieil  to  a  portion  of  the  iietirt  known  a^ 


^    I     i_LX  J^    I     I     I     I  J.  1 


I!     \i 


\    ;l 


'  "■'     '■''       '■■':    '"■'"    lr...niu    .ii,.nii,B    111.    .i>li I    ii....luif.      Th,'    \;ii<ii«    iruiik    ».i<    ^linnil  • 

•'-    ii"lii.it..l        hi    Ihi     iii.il    (liUMii    liii.JMi!    iiiliil.ili..n    .,i,iir<    l.ut    allrr    iiuutir,     lsc.-,.ri.l    lij.r 

11.1   H,liihili..ii    l,.ll.,»,,      .Still, nl.iti. ,11    ..t    thv   ,1,-,,  lit    ill   111.-   ti.M  lA..   liii.s   ~till    1,    t,.ll,,»,-,l   hv    ml. 
"'""        ""     '""'    '''"t-    "1    III'     .li'.i::     111    -li,«i>    111    il,     lanl    Iwi,    iii|.|.,r)    ti.i.ini!-.      I  l-r..in   JjikM- 

t}io  wliite  ereseentie  line,  inhiliition  still  ne<  tirs,  heeause  at  this  jjoint  tl 
l)osftranf.'lioni<-  nerve  tihers  coine  neai-  to  the  siirfaee  iind  tlierefore  a 
stimulated  (Fijr.  (i!ti.  (in  the  other  hand,  atropine  is  a  ih\\\i  whi. 
l)ai'aly/.es  the  postjianv'lionie  tihers.  so  thiit  after  its  application  tu  t' 
lieart  inhihition  c;in  not  he  produced  l»y  stiuuilatiii^  either  the  vat- 
trunk  or  the  wjiite  ereseentie  line.  I'iloc.-irpiiu'  and  muscarine  are  dm 
wliiell   liave  :in   action   exactly  ojiposite  or  ;int,ifronistic  to  that   of  .'iti 


i 


oblongtfj 


Accessory  n 


Spinal 
medulla 
[cord) 

Rami 

communican- 
tes  going  to 
3ymp.  gang, 
{preqanglionkj 
Ansa 

aubdavia 
(Annuiusof 
Vieuisens) ' 

Thoracic 
nerves 


N  I 

Poitganqlionic  fibers 

dre  (Jotted  tttus 


H.VI 

N.X 

Juqular  ganglion  (Cang  of  the  root) 
Depressor  (Tallin  prrssjre  or  slanging  of  heart) 
(Sensory)    ieparate  nerve  n  rabbit  and  opossum 

nodosum  ganglion  (Cang.  of  the  trunk )      '„*";/„*,„, 
Inhibitory  cranial  autonomic  fibers 

Superior  ccrvicol  ganglion 
Descending  sympathetic  fibers  in  cord 
Cervical  vago-SYmpathetic  trunk 


'^Electrodes    (slowing  or  stoppage  of 
Subclavian      heart  Augmentation  in  some 
art     ^  — ■     !v  animals  ) 

>^  Aortic  arch 


first  thoracic  ganglion 
litellate) 


Electrodes 
(AicLeleration,  or 

augmentation  ot  huarfi 


!.i 


...-W  •-•ll  -.. 

i:^ii  -At  !■■  1: 
^1>.,.;.,  l.l„... 
.,1.      I..-., -..I     . 


I     till      I'lr-  T\  .1' 

Hi.!'-        ....  .1  l.t.llM- 

,.ll  V,  ,1,.   .  .-. 


Ill,    1' 


„...l,.  i 


k 


Till.    itiN'I'ltiil     iih     TIM     I  illi  I  I   MliiS 


2-27 


|iii:i-:    tllllt    in,    tlicx     >tinillliil<-    tin-    |ii>slL'iill'.'lii>iiii-    lilurs    uml    ihiiiImi'i-    ii 
Hlnwiiit;  mill  |Mi>i<.ilil\   ill!  I'litVcliliiiir  nt'  till'  lii'nt. 

Ill  till-  iiiiiiiiiiiiiliiiii  Ill-nil  ii  l.'ii'L'i-  iiiiiiiIm'i  III  till'  lilii'i'x  ill  tin'  rittlil 
\imii>  iii'i-\i'  |>iiii'('i'il  iiiiii'il\    til  till'   siiiipjiiiiii'iiliii'   iimli'.    \\liiM('   it    I'liii 

Im'    sllllWII    llislllllliril-llliy    tllilt    I'cillsidtlalpjl'    llll|ssl>    nl'    llrl\l>ll^    ti^^ll^'    l'\i>t 
l»ll    till"    lltlllT    llllllll.     till'    mi'.lt     lllil.iMrity     «\'    III,'     lili.'ls     ill     III,'     I, 'ft     \||!r||> 

luiH'i'i'tl  ti)  till-  iiiiiifiiliiM'iiliiniliir  ImiihIIc,  in   whicli  nUn  ih'imhis  siiiif- 

liiii's  arr  iiliiiiiilaiil    (  |iiiv'i-    Is'  .\»  :ilnMil.\    iii.lii'Mtcd.   tl \|H'iiiiM'iil.il 

ii'siilts  whicli  fiilliiw  still','  ..  "1  illiir  iiciM'  i-aii  lir  rxpljiiiii'il  Ip\  tli  ■ 
lllljlli'lli'i'    ullirll    till-    lli'l     ■     ■■%       Is    nil    r        j    •■■liillliir    stl  IH'tllli-    In    uliirll 

III!'   iiiiijiiiitv   111'   its   (ill  i      ',.<.,;■  .,1  oiiiiiiiliiliiiii   111"  till-    liuhl 

\aitllM  is  likely  to  pi'  ■•  .■\\\<>.  ,  ,  il  w  ■'  ■  .ni;  cil'  tjic  lii'nl,  wiii'iras 
^tiiiiiilatiiiii  III"  llii'    "       ^:i.".-.  i     II'.'  ••  I'kt!'  iiistiliitc  il   I'liiiilitiiiii   "t" 

I'lii  tial  lifiii't-liloi'l. 

Oil  iicciilllil  of  I  ■  'il1ii'  il  1  ■.  Ills  \:  I  II  ,  iiy  111'  iilitiiiiinl  liy  sliiiill- 
laliiii;  till-  vaillls,  ,  •  .1  I  .'  '  . -,  \r.,  !•  .;sv.:||ii'il  tliat  tin'  lirait  iiiilsl 
<ni\\i\ii\  four  ki  nils  I'l  '•■ir  n,  h  ;'i.  i,l  i  t/ns  mrn  i  iiiliiiiis,  niic  t'lir 
1  .ii'li  kiiiil  111'  iiilliiriii'i'    vliii;'   III.'  \.i'.  I      ■    M  (li'M'liip,     'I'Ih'si'  I'mir  iiillii- 

I  iM'i's  lire,  it  will  lie  ii'li  .  '  m.  'li  stiiiitrlli.  tin-  talc  ami  tin- 
IHiipairatiiiii  nl"  tin-  lii-arllu-a;          .  '\\v  ■■^'  ;  .iliilily  nt'  tin-  ranliac  iiiiisrlf 

II  is,  liiiwpvfi',  aliimsf  i-iTtaiiilx'  iiiiiicri'ssary  to  iiiaki'  siirli  an  assiiiiip- 
ti'iii,  for  tlu'  ri'sults  i-aii  In-  rxplaiiu'il  as  niiTi'ly  ili-pcinlciii  upon  ilit'- 
r<'i'i-iit  (1ot!i'i-i's  of  -itiiiiiilatiiiii  III'  till'  saiiii'  kiinl  nl'  lilifi-  aiitl  iipmi  llir 
lAai't  jiart  nl'  tlu'  lii'ail  In  wliirli  llii'  lilicr  inns.  Sniiict lines.  I'ln'  I'X- 
.iiiipli'.  wIh'ii  the  liu'lil  va'_'iis  nnxt'  is  stiiniilati'il  \i'iy  I'l-i'lily.  tlii'i-i'  iii.ix 
'"•  only  a  iliiiiiniitinii   in   lliv   t'nii'c  nt'  tin-  heats  \\itlinnt   any  eliaiiu'c   in 

liiir  rate,  inilieatinir  tliat  tlie  ell'eet  li.is  lieeii  npnii  tlie  ninseiilatnie  nf 
ilie  aiirieiilar  walls  ami  mil  nii  the  simianiieiilar  nmle.  It°  the  sliniulns 
IS  inereaseil  a  little,  then  Imth  an  ellt'eelilini.'  ami  a  slow  iiii;  nl"  heat  nreiii-, 
iiiilieatiii)r  that  the  sliniiiliis  has  nnw  passed  Imth  In  the  aniieiilai'  iiins' 
|ii!.itili-e  direetly  and  tn  the  sinnalirienlai-  nmle. 


<  i| 


The  Sympathetic  Control 

The  efTeet  nl'  the  synipathetie  nerve  nii  the  hear  ia\  he  deserihed  as 
!"im.'  exactly  nppnsite  fn  that  nt'  the  \airus.  The  jiathway  ainiij;  wliieh 
'lie  tihers  nf  this  ncrxe  travel  to  ihe  heart  is  ninre  nr  less  a  ilevioiis  one. 
1  liey  arise  in  the  iiiaiiiiiial  finiii  nerve  cells  in  the  <rray  matter  in  the 
ijiper  thonieie  pnrtinii  nf  the  spinal  enrd.  The  fihers  leave  liy  tho  cor- 
i's|iiini|in!r  spinal  runts  ami  |)ass  liy  the  white  rami  cninmnnieaiites  intn 
'lie  sympathetic  chain,  up  which  they  travel  to  the  .stellate  ami  inferinr 
' '  rvirnl  >;i»"S.'lii>.  Arnund  the  nerve  cells  nf  the  stellate  iiaiiirliini  the 
lil'ifs  end  liy  synapsis,  and  the  axons  nf  the  cells  are  then  eniitinued  mi 


k  ^ 


L»*28 


Tin:  riuci  i.ATiiiN  OP  Tin;  ulckid 


jis  |insiu;;iiiirliiiiiic  lilM'is,  |ir(HT(( I iii<,'  t(i  tlu'  licart  lliii)iij,'li  liriiiiclics  nnu 
iim  otV  t'loiri  till'  slclliitc  "j.iii'.'liiiii   itsfir,  (If  t'rdiii  ilic  n'lsa  suliclavii   or 
iiili'iiiii-  ccrvii-al  -.Mtiu'lidii.     (Ki^',  7(1).     In  i-old-hlimdcd  aiiiiiials,  siicli  a'- 
tlii'  iVoiT,  till'  syiiipatlictit'  lihi'is  run  up  to  tin-  upprr  fUil  of  the  cervical 
sym|)atlictic  ami  join  the  vaijus  ininiciliatciv  alter  it   leaves  the  eranial 
eaxity.     They   then    proeeeil   aloii};   with   this   nerve-    l'oriiiin<;  the   vafr" 
syiiipaliietie     to  the  heart.     The  elTeet  ot'  stimulation  is  shown  in  Fiu'.  71 
The  syiiipathi'lie  nerve  ilitVirs  froiu  the  va};us  in  that  a  imieh  loiijrer  la 
tent   periipil  elapses  liel'iire  its  intluenee  lieeonies  ell'eetive,  and  this  persists 
tor  a  niueh  lonyer  jieriod  after  the  stimulus  is  withdrawn.     If  the  vacus 


A. 


\<K     "1        I't.,.!!!!;"    vlu.nini!    llic-    i  iT.  .  i>    ..„    tli,.    Ii,.,itl„.,|    ,,f    ||.,.    f,,,„    ri'MilliiiK    fr,.iii    stiniiil.ili..M    -■ 
llic      -Mii]    nil.  ti^     iittw-    |ii,i     1,,    ilii.r    i-.Tii  'Ti    mill    till-    V  i«u~    ll.rvi-.       (I'l.ini     l!iiuli.>: 

and  s\aipathi  lie  are  stimnlaled  ;il  the  same  time,  as  Ity  exeitini;  the  vai:" 
sviiipatiielie  ill  the  fro'.',  the  lir^t  elTeet  oliserM'd  is  that  of  the  vairii- 
usually  follow  I'd,  after  reiiio\al  of  the  stimulus,  hy  the  s\  iiipatlietie  el' 
fei't.      If  the  stimulus   is  maiiit.-iineil    foi'  a   loiiir  time,  so  that   the  vjiu'M-- 

lii'ioiiies    lati-x 1.  ese.ipiiiii'ht    will    oeeur  earlier  than   with   pure   vau'U- 

stimulatii.il.  and  au'_'iio'iitalioii  may  lieeome  ai)parent.     The  symp;itlii-tl' 
ililliniiei'    is.    houcxci-.    iie\er   so   stioiii;   .'is   tluit    of   tlip    vatflls.      The   t\'' 
nerves  are  therefore  not  aiil.'ii:oiiistie  in  the  sense  that  the  one  ueutiiili/' 
the  effect  of  the  other:  Imt  when  liotli  are  stiiiiiiliited,  the  hcirt  respmi'! 
first  to  the  vairus  and   later  to  the  sympathetic. 


I  f 


(HAI'TKIi  XXVI 


THK  CONTUCH-  OF  TIIK  ClIfcrLATKiN  (("..iitM) 

THE  NERVE  CONTROL  OF  THE  PERIPHERAL  RESISTANCE 

As  !ilrfa(l\-  fxpljiiiu'd,  tlic  iicivt iilinl   of  tlio  |)('ri|)lu'r<il   ^<•^sistillll•c 

l.ikcs  |)I;h*('  tlii'iiiii.'li  tlic  iictidii  of  vjisiifinisl  riclor  and  vasodilator  iici'vi- 
lilnis  on  (lie  Minsciilatui'r  of  tlic  artci-iolc  walls.  Tlic  vasoconstriftof 
iiripulscs  like  tliosc  in  the  vatrns  of  t!ic  licart  are  tonic,  so  that  wlicn  a 
iii'ivc  containini;  such  tilici's  is  cut,  tlic  coi'i'cspondinp;  Mood  vessels  un- 
ilii't-'o  dilatation  (sec  paue  K!.")),  and  when  theii'  peripheral  ends  arc  stim- 
ulated artificially,  constriction  occurs.  On  the  other  hand,  the  vasodi- 
laliir  impulses  do  not  appear,  at  least  under  ordinary  cireunistaiices.  lo 
\>f  tunic,  so  that  the  cuttinsj:  id"  such  tilici's  does  not  cause  vasoconstriction  ; 
ili'ir  stimulation,  however,  causes  markid  dilatation.  Vasomotor  (iliers 
,iic'  contained  in  most  of  the  cfl'erent  (motor)  nerve  trunks,  and  to 
di  tect  their  presence  the  nerve  must  lie  either  cut  or  stimulated  and  thi' 
rciiidition  of  the  lilood  vessels  (d"  the  innervated  area  ohserved. 


Methods  for  the  Detection  of  Constriction  or  Dilatation 

Several  methods,  varyiii<r  with  the  exa<-t   ai-ca   under  ohservation,  can 

I"'  (isi'd  for  till   ilitirtioii  of  iiism  oust  rid  ion  or  ililnlutioii.     In  nian.\   ea^es 

iisiiiil  iiixiiicliiiii  is  snflicient,  as  in  ilie  w  cli-know  i>  experinienf  of  Claude 

l"(i(ai(l  on  the  Mood  vessels  in  the  ear  of  the  ralil'it  (see  Kiir.  KItii.     W'len 

tills  is  held  with  a   liulit   liehind   it,  and  the  cervical  sympathetic  id"  the 

'■"(■(■espoMdinir  side    is  cut,    maiked    dilatation    will    heeome   evident    and 

'  ■'v-js  will  sprln-;  irdo  view  where  jireviously  there   were   n<MU'  vi>iliie. 

\  i^ual   inspection   is  usually   also  a  satisfai-tory   method  of  demoustrat- 

' '-'  vasodilatation    4ir   constriction    in    exposed    <;laiids,    in    mu<'ous    jtas- 

i-'es  and  in  the  vessels  of  the  skin. 

Another  comparatively  simple  nn-lhod   is  thi'  oliservati<ui  of  lln    h  in- 

rniliirr  of  III!    jinrl,  this   lieini:  i)artii-ulai'ly   useful    when   the   vascular 

■    '  .1  is  one  situated  in  the  petipheral  ]r,\it  i>(  the  hody,  such  as  the  hand 

'    l""t    (see  patfc  '_'(»!»).      When  dilatation   ociMirs  the   tempiTatnic  of  the 

'•     I  rises,  liecausc  the  warnu'r  Mood  fi'om  the  \isceia  Ihius  with  uiciter 

'■loai  throu<;li  the  petipheral   reirions,  where  it   is  cooled  o(T  li\    laclia 

'!(.     When  a  thermometer  is  placed  lietween  the  toi's  of  ;\  dou  or  I'at,  a 

•JJ'.t 


i:hi 


I'lll     1  ||((  I   I    M|(i\     i,K     I  III      lil.i 


llislilll't    I  IM'  ill   li'l|||)c|;i|iiic  XV  il 


lie  nl 


iM'iM'il  u  hi'ii  till'  M'ljitic  nerve  ii(  th, 


I      I 


.•..rres|.o,i,liim    limb    i>  nit       Tlie   ;ii,|,li.Mt  ion   ..f   tliis    ,,iiMei|,le    in   .let.  i 
niimiiir  tli.-  niiiss  iii..v  ,.|Meiii  ,,f  |,i,„„|  |,y  il,,.  ;,,„ ,|  ,,|  he;,!  frjveii  .,fT  fn.i,, 

till-  IiiiIkIs  or    feet    ll.iN  illl(  M(l>    lie,  II   e\|i|;illl.  (I. 

Oilier  Iiietli.p.js  .lepeinl   up .I,sei\  .ilinii  ..f  the  ,H,lllo,r  of  hhmil  fn,„. 

tin    r.ins  „f  ll„    ,..,rl.      \   simple  iipplieiilion  e.f  llii>  ,ii,.tlH„!  eiiii  lie  iise.l   ii, 

" ■•'■^"  "'■  " •"■  "f  ""•  ''ll'l'i'       If  IIk'  lip  of  tl ,•,,•  is  eiit  ..IT,  hiee.li,,... 

"'l<l<'f    "f.lin.iiv    eiiemil>t.iliees    i>    only     \en     sli^rht.    |,„t     if    the    eervieni 

sMlipHthelie     is    elit.     it     I.eeu s    qiiile    m.-|.ke,|.    sln«  in^.    ,l„u  „    „f.Mi,i     nl 

fX.'ll    sl,,ppiM-   eiitiiel\     uheii    the    p.iipJHT.l I    ell, I    ,,f   the    liene    is'stiiiii, 
lilt,'.!        |!y    liiiikiiiM    i,i,.;,sli|-..iii..||ts   ,,f  the    xnjui f   tl,..   ..Iltd.iu     ..r   I.I I 

l'r..iM  il  x.'iii  l,\    this  iii.th,.,!.  III,.  ..Miiii  ,,f  , siri,ti.,ii  or  ,|ili,t;ili,,ii  ..^11 


tube  to  recorder 


oil  enclosed 
by  membrane 


1;     .         ki.jn.  . 


■'"..'I  ill. .Ml       l,l.k....il.l 


'"•    '"ll""-l    .|IMnllt,-,tM..|v,       V.-,s,H|ihit;|ti,.l,    Ills.,    ,.;,i,s..s    eh;,l|.,r..s    in    tl„ 

''""■"'■'"■    "'"    " """'     ''""■■     '!"•     UsIImIIv      >...lltilll|„l|s     II, ,U      1 ,„||i„,r     piii 

:""'"  •'""  " '"••  "'■  "'■■  I'l I  I'nu'ht.  MiM..-      (• .ri {  the  ,,nss,n;  - 

'"   ""    '"''" ""'  ""    ""-  "'   -1   I'^Ht    is  ..Is,.  ..n..,,  „f  viih...   in  f let..,- 

' '    "iKiti.jvs    in    the   ,.,.,|,l,..r   „f   th..    I.I I    v.'ssels,    f„r.    „f   ,..,„,.s,.     th- 

■-'r,.,-,t,.r    II,..    .Iiir..r..|„.,.    ,1,    pr-.ssur,.    l.-.tue-.n    th..    |u „.,n,.|..rs.    tl„ 

-'■'■•'''■'    I'll!-'   !"■  111'.  i.'sis(:iii ir.  .,..1  t,,  th,.  ti,,„, 

'■''"'    •■M"'niM,.|,t;,l     |,iiip,,Ms,     h,,u,v,.r.    ih,.    stall. hir,|     i,„.th„.i     is    tin 

^''"''"  ^"  ""■  ''''"'" ■"■"/''"■■      '■■'•"■  Hms  piirp,..,.  tl .,.,11  .,r  tissue 

'''"■'"'^••''    '"    "    -"••ill..,l    pl,il,XM„„..r;,|,|,    ,.r    x„lu.....    r ,..|..r     th,.    pn, 

'•'•'''■  "'■   "'''■■''    "'"    '■'■  '■l-"l.^    --H    l.v    .•„liMllt,.,ti f  the  ii..e,.,„,,«MMn 

'luij-'nuii    „f    „„e    a,l,,pt,.,l     t„,     ,h,     Li, y    ,|.'|.^.    7L'  Amv     ilierense    ,|. 

"■'■"•'I    h     li'is    u:,-;u.^    II,    II,..    x„|,.l,.,.    „|     ||„.    |.;,.,     ^Mlst     I,,.'   ,lue    e,tll..r    . 


Tin;  iDNTiKH,  (IK  Tin;  ( iii(  i  i.aiius 


2:n 


I'A 


s 
3 


'3 


.III  IrHMCiisc  ill  Miiml  llciwiiii;  into  tiic  \csscls  iicciniso  of  incrciiscd  heart 
iitiiiii  or  to  a  local  vasodilatation-,  anil  vice  versa,  when  shiinUasro  oe- 
.  iirv  We  can  not  tell  t'nnn  the  Milniiie  liaeinjr  itself  whieh  ot"  these 
I  .laiiues  is  really  respnnsilile  tnr  llir  oliseived  alteration,  liiit  we  enn  <1o 
sii  li\  .-jiiiiiiltaiieously  oliserviiiH;  the  mean  -ii'tei'ial  lilood  pressure.  If  this 
tills  when  the  volunie  decreases,  it  nieans  that  th<'  volume  of  hlood  ilow- 
iii'_'  to  the  part  must  have  liccunie  diminished.  If.  nn  the  other  haiul  the 
l.liM.d  pressuii-  icmains  constant  or  rises  uliile  the  xolmiie  decreases,  it 
iiuans  that  the  lilood  vessels  have  locilly  constricted. 

Methods  for  the  Det«etion  of  Vasomotor  Fibers  in  Nerve  Trunks 

If  we  wiJi  to  lind  out  throu'.'h  which  ncr\c  tiunks  a  iri\en  vascular 
;ina  is  supplH-d  with  \  asoconsi  nctor  or  vasodilator  imiiulses,  we  should 

liriiceed  liy  the  use  of  oi f  the  alioxe  dcscrilied  nn-thods  to  olisorve  the 

ifTcct  priMliiced  on  the  \cssels  li\  .iitlin!.'  the  ncr\c  and  then  liy  stimil- 
iiilini:  the  peripheral  end  of  the  ciii  ncrvi'  As  a  result  of  such  ohserva- 
;i.ins   it    has   heen    found    that    the    \asoimttor    liliers    nv    frei|uently    dis- 

•riliuted   so   that    those   havinir   a    vas iisirictimr   action    are    collecteil 

iiiainly  in  on<'  nerve  trunk  and  those  haviiiir  a  dilaiiMs:  action  in  another; 
111  some  ner\c  trunks,  however,  the  relative  inimlieis  of  the  opposinir 
lil.ci-s  are  about  ei|Ual.  Nerves  conlaiiiiiii;  a  iire;i*  prepoiuierance  ol'  \  aso- 
•  MiisirM-tor  fihers  are  the  y:rc,it  splanchnic  and  the  ci'rv  i<-al  symjiathelic ; 
.,ii,|  tli+»se  coiitaiiiiiic:  a  irrcat  preponderance  of  vasodilntor  are  the  chorda 
i^iiipani  nerve  to  the  suhmaxillar.v  i:lan<l  ami  the  ncrvi  eriu'cntes  to  the 
.\ii'rnal  tretdtalia 

It   must  \>o  elearlv    umlerst I   that,  althoiiirh    one    kind    of   vasomotor 

I'liir  may  jirepondcrate  in  one  of  tli»^c  nerves,   vet  the  opposite  kiml   is 

Iso  present.     In  the  cervical   s\  in|»Hthetic,  foi-  example,   some   vasodila- 

•..1    tillers  cNtendiii'.r  to  the  hlood  vessels  of  the  mucous  iiiemlirane  of  the 

■  sc  and  cheeks  can  readilv  lie  demoiistiated.  as  shown  liv  the  llushiiiir 
.   these  parts  when  the  peri|>licral  end  ot    the  nerve  is  slimulalcil :  and 

■iilarlv,   ev-'u    in    the    >rrcal    splanchnii-    nerve    itself,    vasodilator    tiU-rs 

■  plviiii:  the  suprareuid  cipsulc  can  i|uiie  readii.v  Im>  made  nul.     When 

vaso<'onstrietor  lihcrs  t;rca1ly  prcponder.tte  ovei-  the  vasodilator,  'he 
•  <'t  of  the  latter  ma.v  he  demoiistiated  l.v  taUin<_'-  adv.intajze  of  the  l.iet 
.1    I  niiilii.riui     par.il,v/es    the    v  asoconslriclor    but    not    the    vasodilator 

■  rs,  so  that  after  its  administration  siimuhition  of  the  irieat   splaneti 

nerve    jrives    rise    to    a    vasodilatation    instead    oT    a    v  asoionsf  riel  ion 

■  prtsenee  of  \  asoeoiistrictor  tiliers  in  the  so-called  vasodilator  nerves 
Mida    tyiiipani   and    iierv  i   eritreiitcsi    has   not    however,    heen   demon- 

,ted. 

\    L'ood  c\aniple  of  a   nerve  'rnnk   containinir   aiioiit    an   eipial    ailmix 


Itoi 


Tin:  riK(  Ti.ATiov  OF  Tin: 


III.MOj) 


diri'  of  hotli  kinds 


•  •r  vasoiiiiitiir  /ilxTs  is  tlii 


sciatic.     If  tlic  liiml  liiul 


>  <ii 


.••  .l.-ir  ,s  place!  ...  a  plctlnsino^r,,,,,!,  ,„„i  ..i„„iifan....uslv  a  m-o.-.l  of  f|„ 
'"•''•'"  "'X''";.!  M.,0.1  pressure  taken,  it  uill  he  foun.!  on  cutting,  the  sciali, 
nerve  that  fho  volume  of  the  Mini,  increases,  whereas  the  I.loo.l  pressure 
-vn.ains  ,vract>cally  constant.  I'.efore  placin«  the  liml,  in  the  plothv.sm,, 
«n.|.h.  the  inus..les  must  of  course  he  paralvze.l  hv  n.ean.s  of  curare 
otherxvi.sc    n.usc-ular    contractions    woul.l    confuse    the    result.      If  th.' 

peripheral  en.!  of  tl ut  nerve  is  non   .stimulate.!,  va.soeoiistriction  vili 

'7"'''"  ' '"^'-'v.'-l.     So  far.  then,  the  results  .Icmonstrate  the  presence 

of  vasoconstrictor  nerve  lihers  alone. 

To  ,!einonstrat.'  the  presen. f  vasodilators  a  difTcrent  procedure  is 

•u'.M-ssary.  This  is  ha.sc.I  on  the  foliowinj,'  fads:  (I)  The  vas-Klilatm 
liervo  hhcrs  detrrnerate  nior,.  shmly  than  the  va.soconstridor ;  (2i  thc^ 
.■ire  less  depressed  in  their  e.xcitahility  hy  coolini;  the  nerve;  and  (:{)  thrv 
are  more  sensitive  to  weak   slow   faradie  stimulation  than  the  vasocoii 

strictor  fillers,     A. rdinudy.   if  we  cut   the  .sciatic  nerve  two  or  three 

days  liefore  the  actual  esperiment.  and  then,  while  ohserviin^  the  volmii, 

"•■  ""'  '""•'•  •"•"• 1  '"  stimulate  the  half-dctreneratc.l  nerve  with  feehl. 

electric  stiiiinli  of  slow  frc,|Ucn,.y  we  shall  usually  ohserve  a  dilatation 

of  the  limh  in.stcad  of strictioi,;  and  even  if  we  cool  a  .stretch  of  a 

freshly  cut  nerve  hefore  applyin-  the  stimulus,  the  same  result  will 
ot'tcii  he  (il)taiiied. 

The  Origin  of  Vasomotor  Nerve  Fibers 

Havin.,^  seen  how  the  iiiTsence  of  va.son.otor  fihers  mav  1 etctcd  in 

'"'.''''''"■,'"'    ''"''''•'■   "■■   "'"^'    ""«•    I"''"-"-l    to   trace   them    hack    to    ,h..i, 

..nun,  from  the  central  nervous  sy.stem.     The  method  for  d. this  co„ 

sists.  in  .,,n,.ral.  m  ohservin-,.  the  etTcct  on  tlie  hloo,|  vessels  pr..duccd  l.^ 
'■uttin^.  or  st.mnlati.ifr  the  various  nerve  roots  throuirh  wliich  the  IIIm  ,'. 
ninrht  pass  on  their  way  to  the  nerve  trunks.  As  a  result  of  such  ohscr 
v"t...M.s  1,  has  heen   fouii.I  that  all  of  the  rn.ocunslrirlor  /Ihns  emaiial. 

''■"'"  ;'"■  ^'""' '■■'  "'  '•"•  '•'•^'i""  '"■•"cen  the  level  of  the  .second  thoraci,- 

'"'<l  "'="  "f  the  sivond  ..r  third  lumhar  spinal  roots,  hut  from  .lowher. 
else  m  the  cerchrospinal  axis.  Section  of  the  si.inal  cord  helow  Hie  lev.  ^ 
of  tl...  s,...on,|  lumhar  spinal  r..ots  i.n.duc.N  n..  .■han-.'  in  th..  v.,luni..  n. 
""•  I'll'. I  limh,  provi.l...!  tl,..  muscles  he  tlnn-.-uirhlv  curari.e.l    nor  .1...- 

'"'""''"" '■  ""'  l""<'i-  ''"'l  "f  11"'  <Mit  spinal  c.r.l  hav..  any 'eir.vf      P 

;'"'  '•■'"'  '""  " ■'••''•  "••  »'"•  <ii.^'  two  luinhai.  spinal ts  arc"  .stiinulat,  •: 

liowev.'r,  .•\i.!..n..c  of  vas.M...nstricti..n  will  lie  ohtained. 

The   .•.■stri..ti..n   of  th..  ..ri-in   .,f  va.s nstrict.ir  (ihers   t..  the   -ih.n 

""■"" ''   '■'■-''""^  "•'  "!•'  si'iiii'l    '•'"•'I   in.li.-atcs  that    in    i.r......,..lii„.   ,, 

th..    mi.xe,!   nerve   trunks   lh..y    must    travl    aloni:   s| iai    nerve   .."tl 


Jt  ! 


Tin:  ((isiKoi.  UK  Tin;  i  ik(  i  r.xTiciN 


2:5:1 


'i'licsf  iMT  lH'ovidcd  l)\  til.'  Nyiii|i;illi(|ic  rli;iiii  .iiid  its  IhmiicIics  (Fin.  'i'iSi. 
Tin'  vasopoiistrictoi'  (ilxis  in  tlic  jiiiliiior  spiiiiil  niots  leave  tlio  Iiiftcr 
l.y  way  of  flic  coiTcspoiiiliiiyr  uliit,.  i;inii  coiiinimiicaiitis,  mikI  jkiss  into 
tlic  nci<;lil)orin«;  syni|)aliii'lic  cliain.  almij;  wiiii'li  tliry  citlicr  ascend  or 
descend,  accordiiitr  to  tlicir  nltiniate  destination.  In  llieir  course  they 
(•(PHie  into  contact  with  tin-  syin|iallictic  ifan;rli:i,  tlifoiisrii  oin-  or  two  of 
uhich  tlicy  may  i)ass  «itlionl  any  cliansrc,  lint  \iltiniatelv  each  liluT  ar- 


n\cs  at  some   uanirlioM.   in    uliicli   it   terminatt 
.iioinid  one  of  the  i.'an«lionic  ncr'\c  cells,     'j'li 


s  liy    t'i)iiniin,'  a   synajisis 


e  a\i>ii   uf  this  nerve  ct 


ilicii   cd'.liniles   iIh 


|.\     I 


ic   iiiar'c  t    '_'i;i\    laipius  i-.«!j|iiinnican 


111  the  spinal  nerve  Iic.mhmI  the  union  ni'  its  antciioi-  an< 


I     )>MHt 


II 

s   Hack 
crior  loots. 


I'p  to  the  iioiiit  where  the  lihcr  forms  n  synapsis  with  a  jjanylionic  nerve 

the  |irc'_'aiiu'liiinic  lihcr.      |{cynii<| 


it  is  mcdullated  an 


i|    IS    Klliiwn    ;i' 


I'cll.    it    is    iii>niiie(lullaicd    aii<l    is    kimv,  n    as    jiost'_'aii;;lion 


1 


Til.-  .Aiicl   ^aiiylh 
•    .    .•.•II   .;:ii    !,■   .1.1, 


II   ill   wlii.'li  .1    i^ix.ri   \: 


iii-tri.t.H    lil.,r   I.. 


'illlirrti-.l    with 


■.!   l,v    II,. 


Iii.-..I|M.'    liii'tli    .1    .it    l.iiiiuliv.       1 :il    ;n,|ili.-:it 


111.     -.uiulimi   of   il    tti'ik   s.illlli.ili    i.r    tliis   ,lii|M     ,  I 


thill    :i   stiiiiiiiiis   .'i|i{iii.'il    In    II 


|i|i| 
I'.r  .■.III  1    |.:n!.|i  /.■>  tl.i-  svii:i|.tic 


|iii'L:;iii:;li.iiii.     lil.. 


n-^i-t     [.iii.lui- 


It- 


I  111- I.     Mii|.|i.>sc.  f.M-  .xiiiiiiii.'.  t|i:,t  :,   v:im.,-..ti«ti  i.-t.ii   lil..r  liiis  1  ..en   f..UM.I  l.>   Il».  -liiiiiihi 
li"!i    iii.'thii.l   1.1   tr:i\cl   tlir.m«li   s,-\ri:il    ;;:iii-Iki,   ;iii.|    >..■   wl.li   1..   ,1,'ti'i  nu.i,.    iit    Hin.-h   i.f 


III'  N_\  (t:i|isis  MIS  :    we 


.1,,  ,     li- 


:.|.|.| 


lhi>  stiniiilii 


;it    .-1     |i..ilit    .•illl  !;il    In 


;;:in','ll;i   iill.r   iMHiitiii-  .mil   i.T   th.Mii   in   1  ijni    with   tlir   iih-i.i  in.'   s.iliil  ii.ii.      Il    ih. 


:.l|||ir:ili.ill   lit'    I  h.'   ilrii^   I. 


^.i\rll    ;;;ill;;ti.in     in    I". 


Illl   :ilti'r:ilii.ii    in    ih. 


.  n-,.i    iiriiilu 1    li 


y    stillllll;il  1..11,    tllrll    «.■    klii.w     tli:il    I  li. 


Ill    n.il    I.I 


itiiiii   ill   tliiit    t'aiiylii'ii.   mill   we   i.i..it,.I    in    il 


111'    >;illir    u;i\     till    Wr    li: 


■viiniitl.' 
l...-il.'.l 


::oi^'liiiii   ill  wliiih  syii:i|.»is  iir.i.is.      h    i~   iiii|.iii 


piiitjiiii    ti 


iiii.iiil.i T   lli;il    till 


iiulii.iiir  va.mi<'iiiistrii'l.ir  lil.ns 


II  .'1  yi;i>    iiiiiiiis 


iit^  .).. 


.  i::iM:;lii.iiii'   til 

111.'  riiiiii  at   a  liiijli 


lint    .■..III,'    t'r.illl    III, 


if   1 1.1 


s|ii.|i.|iim 
IT     li'M'l. 


ii  i.  lit.  I  It  r 


Tli,'  al'i 
111.'  iiiili'ii.ir  ami 


iii|ili.iii  a|i|ilii's  1.1  III.'  \:i«.ii  iiH~tii.l..i   lil., 


11.111    til'iis   I'.iinin:;    tli|iiii:;li 


■iliii-  til  till'  M'.M'U  iiT 


1     [iiisti 


lliil'    i'\lli'liiilii's.    tlii'M'    Illl    the    f. 


iiu'i    :i  t  i>iii^    I  111 


111   Illl'  I'liintli  tliiiniiii-  t..  till'  ti'iith;  :iiiil  ih. 


r.ir  Ih.'  latt.i.  ri..iii  III,'  |., 


-I   III. 


■  I    till'   first    tllll'i-   l,lllll>.'ll    lirlAi'    |. 


Thi-  ii'll  slati 


111   f'.i    iLi'   lil.iis  ti.  ihi'   I'.iii'   liiiil. 


Illl'  stillati'  ^aiinliiiii.  I    l.ir   tl..-  hiinl  liml.i   in   tli,'  l,i 

il   ^aii^lia   lit'  the   alulnniinal   >\iii|i.'itlii'tir  I'haiii. 


Iiiiiil 


liiil.Mr  a.i 


I    lii«t    Iv 


iinstrlctiir   lllu'is  t"   Il 


ssi'ls   i.t'   till'   h.'i.l   nil. I 


iiiii'ulial    .111'- 


lit    I'liinsi.,    tlii'i-r    I 


;;    111 


I'l. 


iiuii'.iiii-  tllii'i.s  may  mn.     Tin.   lil..i>  t.i  tin.  |.|..  ,| 


«|iin;il    niMi'    al-m;,'    whiih    tin-    |..,vt 


I'ls  ..I'   Ihi'   ill  a. I    h' 


■i| 


1.11.1  t..  the   I'liniih  m    lillh   th. 


iiiiiiiiiiiiiranti'H  intii  tin 


nil.  lil'  ih: 


111,  1111  \\ 


.1    |.ass   1,^    Ih. 
In.  h  th,  \    luii 


!ilr 


:.'itii;liini.  the  ansa   siilulavii,  aii.l   lli,'   inf.  11. .1    ..  im.-.iI   .4'iii:;li,,n.   ilu 


|.   till 

.till'.;    whit,' 
|.a--inu'  tlir..i|n|i  ||.,. 


I'l'rviciil   sympathi'tii'   tn   llie   >ii|.i'i 
Till'   juiNty.'inyliimii.   tilins     .ii    ha^iii^    Ihi 

:il.iii>;  the  niitir  walls  i,f  tlie  M 1   xissi'N 

laMii'iiiistiii'lnrs  til   the  iilii|..iiiiiial 


i'i\ii'al    Laiiuliini.    wln'ii    Ih. 
inuliiin    IraM'l    t.i   th, 


asil'llilill; 

.11    slatiiii 


ili'slinali 


"Is  iif   uliiili  ci.nn'  ..fr   fi.ini   iln-   1, 


I'la    .'111'   call  11 
uri    si'Mli    Ih.il.'i.',. 


.1    l.v    Ih.'   s;  hiu.  h) 


i.l    till 


I'llihisl    liiliil.; 


2U 


Tin:  (  11(11  i.Aiiu.v  (IK  Tin:  iii.(niii 


riHits.      Thr   Ihi.iM.i.-    IIImik   |.:ir.s   il..«ii    tlj.>   ■..VTn|.:.tlictii-   iliiiiii.   «l,i,li    lli.v   Irtivc   l.v   tli. 


^K'llt     .S|>lllllrllllii'     Ih'Mi 


Tin'    llirillilll'    liln 


iH   Cniiii    till'   l('«!<i'r  iir  !iliil"iiiiii:i|   !«|i|iuirliiii 


if  If 


iiiTVCM.  As  |.r.';;.'iiulii.i,i,-  til.i'is.  lli..uf..i.'.  Mii's.'  lil.rrs  :iri'  .•iiriinl  \,y  ||„.  ^rnil.'r  nn.i 
li>Mr  .'■i.laiii'liiiir  iHTvi's  inli.  tlir  .•ili.li.iiii'ii.  uli.i.'  Ih.'  f.iriiKT  i'imih's  iiit.i  iliiw  r«'1iili„i, 
«lii|i  uitli  111,'  '<ii|.i:imiiil  ul.iii.N.  yiviii-  „fT  a  liiiiiiili  to  til.'  Kupraii'iml  u'lmtlli"".     "Hi. 

'"■'''"  "■■•«•  "'■  ""'  '"''»<■  i"  "•"iiliiHi''il  1.11  til  till-  Millar  pl.-xtiH,  in  tlii'  vailniii.  cnnulia  "f 

wliiih   must  iiC  111,.  |,r,'uiiii;;li,.iii.-  lil.,i>  ,.|i,|   |,v  sviiapsJ!.,  tlio  |,ii>li,.aiiyli„i,l,'  filiors  tin.. 

I ■ 'i">-'  »'"»«  III''  ''I I   Ms-'il"  t,i  III,'  v,'«s.ls  „f  til.'  al>.l,.iiiliiHl   \ij.,-.'ra.      (S<v  iiU. 

]<tit:r   S7|l|. 

ViiKodiliilor  jiht  rs  Imvc  .-i  iiiiu-.'  vjiiic.l  (irijriii  tliaii  viis iistricf.ir,  ii!i,l 

flifv    mil   nil    ('iitiifly   (lilTi'icMl    .•.mi-sf.      Viiso.liliil.ir    iiiiiiulscs    iniiy    l„. 
tr.iiisiiiitltMl  liy  tilicis  jifisiiijr  t'nnn  pni.'tically  iiiiy  Icvrl  of  flip  (•cfclin, 
spinal   axis,   ii.it   unly    iiy   tlic   niiitiir   nidts,  liiit   iiy   tli.'  st-iis.iry   lis  w.'ll 
Tims,  Ihcy   pass  out   iif  tin-  spinal  cnid   in   (lie  imslcri.ir  sacial   r.mis  1.. 
•Mifcr  tlif  nerves  of  the  liind  jimlis,  as  lias  lieen  (leiimiistrale.i  liy  <»l»sei-\ 

iiijT  an   in.Teaso   in  the  v.ilui if  the   .•iiniii/.ed   liiiili  (liiriiifj  eleetiienl 

stiiimlati.m  «if  the  exp.ise.!  m.itlets.  The  apparent  ineonsistciiey  of  thes. 
iiliservatinns  with  the  wellkmiwii  law  eiineerniiifj  the  direetion  of  tli. 
iinpiilses  e.uitaineil  in  the  iMistcrinr  spinal  roots  is  explaiiieil  by  assuin 
iii<i  that  the  ilihitor  impulses  are  tr;iiisiiiitte<l  al.injr  the  ordinary  seiis.n  \ 
tillers,  since  then'  .ir.-  no  :'lTer('iit  liliers  in  these  roots.  They  arc  iiiipnl 
ses  whi.-h  ^r.,  airainst  the  .inliiiary  stream  (antidromiel.  In  support  i.i 
this  explanation  it  is  of  iniportanee  to  note  that  at  their  teriiiiiiatinn 
near  the  skin   many  sensory   liliers  split   into  several   liriiiiehes,  some  ,.' 

which    run   tn   IiIimkI    v.-sncIs.   an.l   ntlii'rs   to   r ptor  .irfrans    (pa^e   7!IT 

.•^timiilalion  of  the  latter  liranches  may  cause  dilatati.iii  of  the  lin-al  I1I0...I 
vcxsels  nearliy,  indieatiiij:  that   impulses  must  lie  traiismitfod  up  to  tli. 

I"  "it  at  which  the  lir; Iiini.' occurs  and  then  down  the  vascular  IiraiH-l. 

I  result  lieih-r  olit.iineil  ev.'ii  after  the  main  trunk  of  the  nerve  hit- 
'  ■  •    1  .'lit  ali.A f  the  di\  isiiin, 

*'"  ""■  '•' '  »■•->'•!-  "I'  'I'c  init.ii..r  .■sli.'iiiilv.  tli.'  v!is,..lilal„r  impiilsi's  arc  siiiiihiil 

i..-.i,-.niitl.',|  tliicimli  tl,.'  |.,.st..,i,.i  -.[.iiKil  r  ,.t-  ,.f  ll„.  |„H,.r  ,-,.ivi,.i,|  r,'-i,.n  ,.f  lli,.  s|.l.; 
••.1.1.  TIm'  v:iM..|il!il..i  (il.,'i.  I.,  ||„-  .■.l„i,.iiiiii.il  NisiMTii  at,'  liaiisihllti',1  with  the  .splani-lii. 
'..lA,-,  l.iit  lliii  n,:n  i,l>..  I...  .U:\u.\  Cimiii  iI,,.  p.,sl,'ii„i-  »|,i„:,l  r,".ls.  f„r  it  has  1"  ■ 
f 'I'll. I   Dial  «ti,„,il.iti..M  ..r  p,.>t,r„„    ,,„t.s  ill  .1,,'  >plaii,-lii,i,-  area -■aus.'s  ,lilalati..n   in  t' 

'"''■"" llii.»li-i.      Va»,..|ilal..r    ..l.,is   ar,-   ah,.   ,-..„taiii,-,l    in    tl raiiial    imtv.'S,    ]■■ 

li.'.ilarli  tl,,'  M.,,.|„l,  an.l  tli,.  aiiilli.  l„'i,iM  .JiMril.iif,-,!  in  the  fi.inn'r  iieiA,'  t„  th,'  :. 
t,'ri„r  p,,iii,,a  ,,f   i|„.  .,,„:,„,.  aa.l   tli.-  .alivai^    L:la»,|s.  aihl   in   tin'  latl,T  t,.   the   p..st,L 

';""'""  "'■  ""■  ''■"-'"■  '""I  ""■   Ni.e-.i-   iMl.ian,'  ..|-  tli,    tl...  I    ..(  Il„.  „i„„tl,.      The  ^■.. 

Mal..i    til.,.,.  I,.,   (!„■  ,iiii,-.„i.  1,1,  ail. , an,    ..(  tj„.   ;„vi,|,.  ,,f  ||„.  .-h,  ,.k>  aa,l   naii'S  hax,.  H 

"""-■   '"    " iw,.,l   sMi,|.atl„ti,-.   l.,-ii,-   ,i|v|ril.iil,.,l  to  th,.  I ..I'arial  re-i..ii  in  ' 

l.iaM,h,.s  .,(■  (|„.   til'lh   ,i.ni;al    ii.n.'. 

Thi'i,'  IV  ,.\i,l,.,„,    I,,  .!„,„    ||,M    il,.'  vaM„|,|al,,i    lil„iH,   lik.'  tl„.   va, .iiM  i  i,'t..r.   l^.  ■ 

c„nn,.,l,.,l    l.v    ^Miai'-I-   uiil,    a,-,\,-  ,-,ll«   s.,ri„'ul, i„   their  ,..Mir».'       In   the   .-aw   .■(   • 

'■ '''•''"'    ''l-'i"    1"    '!.'■   .Ii.'i.la    lM"|;,i.i    a„.|    iMTii    ,.rii;,'nl,'j,.    m.h,    ,.,.||    ,tati,.nH    I 

■■'""    'I'-^ol.v    .1.1. -Ii,al...|    in    111,     hiliis   ..1     l!,.-   Mil.iuaNillarN     ulaa.l    ,a    Ih,.    (...inei    ,.. 


TIIK    KINTKIII.   (IF    Tin;   CIKCI  I.ATION 


235 


11. 1    ill    till'    liv|iiiyjiiilrii'    [.IfMis    xitimti'il    i>ii    tlif    iici  k    <•(    llii'    liludilcr    in    llir    liittcr. 

Ii   it  tliiTffiirc  (■'.fiiiitiiiily  immiiiu'.l  lli:il,  ,'iI|Ii<m||;Ii  ii.il   i i;;iii/.iilil<'  liy  lii»tcilipyir  iiictliixls, 

,  !■  h  tiTniiiial  It'll  sliitidiis  iiiiist  til^n  i'xi«t  In  rlnw  iisonriiitinn  witli  nil  lilnml  vi'hsoIh 
'..  wliiili  llir  Viixmliliifiir  tiU'is  iiiii,  Wlii'lliir  cir  mil  sinli  |ifi  i|'li('iiil  irll  statimis  oxi»t. 
'Ii.ic  is  a   niiiikcii  ilifVoiiiiic  hi'tw^'i'ii   llic  iniirw  m'  vnM'Mlilntoi    ami   nf   va!»<MM)nsfrirtor 


The  Vasomotor  Nerve  Centers 

Our   iK'Xl    |ii'ti|il('iii   in   til    tiiiff   llicsc    lilins    fiirl 


I 


icr  into  tlif  (■('iitnil 
I  .ivdiis  s.vslciii,  anil  liiiil  tin-  liiciitinii  iind  stinl.v  tin-  rlianicfi'i-istics  of 
III.'  iicrvf  wntcrs  froin  wliich  tlu-y  aif  ilcrivcd,  Wr  must  iiostnlafc  tlif 
.  \isti'ii('c  of  lioth  vasociinstricliii'  ami  vasinlilatnr  i-i-iitcrs,  Init  siiu-c  tlicrc 
!-  Ml)  adequate  evideiiee  af  llie  pieseiit  time  wliieli  eiuiiiles  us  to  loento 
till'  latter,  we  must  ciinfiiu'  uur  attention  to  tlie  vasoeonstrietor  centers, 
Tlii'se  exist  af  two  levels  in  tlie  i-ereliros|iinal  axis:  (1)  in  tlio  «ray  mal- 
tr  lit"  the  spinal  ninl,  and  C.M  in  the  ^'la.v  matter  of  the  meilulhi 
'  liloiisfatH. 

Tlie  spinal,  or  as  tliey  are  id'ten  ealled,  the  siilisidiary  vasoeonstrietor 
.■I  uti'is,  are  represented  liy  eertaiii  cells  of  the  lateral  horn  of  pray  tiiat- 
irr  in  the  thoracic  poition  of  the  spinal  cord,  from  which  the  prejran- 
L.'lMiiiic   vasiicoiistrictor   fillers  aliii\c   descrilied   are   derived.      The  exact 

!.ii':itiiin  of  the  nerve  cells posini:  the  chief  centers  in  the  medulla  has 

ill  ;is  yet  lieen  definitely  made  out;  they  undoulitedly  lie  near  those  of 
!li.'  vMfrus  center  (see  Kansoni.  The  asons  of  the  medullary  cells  de- 
-iiitl  ill  the  lafi-ral  columns  of  the  spinal  cord  to  end  liy  synapses 
■ii'iuiid  the  cells  of  the  sulisidiiiry   vasoconstrictor  center  in   the  lateral 

!"iMs. 

Till'  cxiierimeiital  evidence  uhich  indicates  the  existciice  of  chief  and 

-  .'-iiiiary  centers  is  <|iiite  definit.'.     Tims,  if  the  spina!  emd  is  cut  at  the 

'      rr  cervical    reirion    (lielow   the   plireiiic   nuclei,   so  as   not    to   interfere 

'li  the  iiiiivcmeiits  of  the  iliapliiairm   ,  the  arterial  Mood  pressure  falls 

'iiimlly.  Iieeailse  the  pathway  conncctiiii;  Ihe  two  centers  is  liroken. 

I- several  days,  howevei-,  the  lilooij  pressure  will  "radually  rise  au'ain 

'lei-  this  has  oeciirrcd.  the  spiii;il  cord  is  dcst  mv  ed  liy  pus|iiii>;  »  wire 

II   the   verteliral   canal,   the   arterial    Mood    pi'essiire   will    aaain    fall. 

I'.itinir  that  the  vascular  tone  which   had  liecii   reaei|uii'ed  after  sec- 

III'  llie  palhwa.v    lietweeii  tile  main  and  the  sulisidiary  ecntcis  iiiiist 

•    lieen   liroii'^lit   alioiit    liv    I  lie  development    in   the  sulisidiary   centers 

II  iMi|e[K'ndciit   power  of  rcllex  tonic  action.     This  I'xpeiimcnt   tlieie 

ileiiionstrates  that  in  the  intact  animal  the  sulisidiarv   centers  do  mil 

heiiisejve^  discharire  tonic  impulses.     In  other  words,  the  suhsidiarv 

■■■i>   ordinarily    serve    merely   as    tiansfi'i-   stations    fi-r   the    tonic    im 

^  ciiiniiiir  from   the   chief   lenler.   Iiut    when    these    iiiipnises    no    loli!.'el 


AJ^ 


L'.it; 


Tin:  ciiii  I  I  Alios-  OK    iiii;  ni.utiM 


.•Iili\r.  111. 11  ;i  lijltiirtii  iliiriiiMnt  pnwri-  i.l'  Idiiii-  ti'Mivilv  lifcomcs  i|i-vi! 
ii|)fi|  ill  llic  Milisiili.iry  rciiliMs. 

Independent  Tonicity  of  Blood  Vessels 

llv.'ii    jifl.r   ,-nin|.l,tr    .liM-..iiiMrt  i.Mi    ,,|-   tli,'   spi,,;,!    ,.,,n|    fn.m    til.'    hi I 

v.-s.U.  .1-   h.v    nitiinir  «\-  III..  >|,|j,M,.ji„i,.   ncivf   I,,  tli.'  ;ili,|.> ,•  ,,i.l., 

"'"I   "I'  'li'il    l"'iti..ii  .,1    til,.  l.,u,T  spiiiMl   .•..nl   IV..1I1   «lii..li   til,.  Iil„.r>t  t.. 

""■    ''''"'    'i'"''   "'■i-^''.    'Ii''   'li^< iifH.'.l    1.1....,!    vcsM'Is.    ,illli..iis;l,    i,t    liiv: 

'"Mi|.l.l,l\  ,lil,il,.,|.  ni.iy  Ii,|..|'  i..,i.M|iiiir  ,111  iii.l.'p.Mi.l.'iit  t..ii..  ..r  tluMi 
"^^M.  ili.li,.;,li,,i;  lli.-rrl',,,-,..  tliflf  Ihry  l.mst  |m,ss..ss  so,,,,.  ,i,..l,„„„is.Mll;i. 
iM.'..|,;,iMsiii  ulii,-|,  ,.,ni  ,•„•(  i,i.|,.p..n.l,.|itly  ,.f  111..  M.-rvc  ,'..|iI.ts,  m„l  wlii.l 
iri.i.v  1...  siiMiiil;it..,|  to  ii,-ii\iiy  l.y  tli,.  pivM'ii.-,.  .,f  h„nn(>nes  in  tlio  I.Io.mI 
'I'll''  li'-nnnn..  «^is  :it  ..ii,.  tiin,.  tli,,ii-l,t   I..  I pi,i,.pl,,-i,„.    ,  y,.,.   p,.,,,,..  74.-, 

Kpi,i.pl,iiii,.  .•..iit,-.,|   is  iii,li..;,t.'.|   ill  til,.  ,.tr,.,.|    pr.Mlii,-,',!  up..,,  i„i,.,i,v 

''' '     l"'"^^"'"     I'.V    Milliul.lti.,11     .,)•    til..    -IVMl     splillK-lllli..     „..|v,..        CHIvflll 

iiiNiK>is  ..f  til.,  .•mv...  sli.iwii   i„   l-\r:,  o,,    ^|,„„^^  ,,,.„   ,,^^,  ^.j^^.  j^  ,^^_j|^   .^^ 

' ''"'■•  ■■""'  ''••I'ly..,!:   tl,;it    is.  til.,  cun,.  i„..ii,its  i,ii„i,'.lii,t,.ly,   tlicM    ll;it 

I'H-  -lit  ,1  litil...  ,,,1.1  ,li,.„  .-isMM.i.'s  ;,  fiiitlicr  lis.-.     This  i\,'hn,'.\  n.sp..i,., 

S...IIIS  1,1  ,|..p,.n,|  iip,„|  th,.  ,.\..,,.|i,,„  of,.pi,„.p||,.i,„.  into  til,.  1,1 1.  r,„.  , 

■i.Ms  II,,,   .„.,., II-  «li,ii   III,.  s,ip,-,.,,-,.,i„|   \,.iiis  ;,,•,.  ,„.rlii,|...l.  aiul  is  iiili,'],  .1, 
I.-I,v,.|    li.\     t,.h,p.,|-;i,il\     .-l.-mipi,,,.'    II;..    slipi;,,,.,iiil    v.'i,ls    .,,1    111,.    s;i„„.    si,|. 
lis  tli:il   ,11,  \\lii.-li  il,..  sphin.-l,,,;,.  ,„.,.v.'  is  stiiii,iliit,-,l.     It  l,;is  liccii  stiil..! 
l.y   .■.•rtaii,   ..liM.rv.Ts   tlinl,  ,-,ri,.|-  ,,....|„si„i,   „r  tli,-  .•i.hviijil    v.-ilis,   tll.T..   i- 

.•I   .l.iunuai-.l    t..ii.l,.n.-y    .,r   ,|„.    I,| |    pivssiin..    uhi,.],    1,.,u,.v.m-   .l,.v,-l.,p 

Milll    ••Ntl-.-,,,..    s|.,„,|..ss;    ,.,11.1    ihai    ;,   .|isti,i,-t    .■l..vati.„i    .,f   I,l,.,„l    pivssiii 

InlL.ns  til,.  i-,,„„v.-il  .,1  a  .-laiiip  t..i„p.„-ai-ilx    p!.- 1  „„  tli,.  a.Ir.'nal  v,'i,i. 

''"''"*  '■'•^'-  '"  I"'"' '•'■'!    i''  ••"■  M'l.-iM.-lnii.-   I,.., -v..  is  sliiiiulat.'.l   .|iini,._r  ,|:, 

•  ..•..|iisi,,ii  .,!•  i|„.  v.ins.     It   anisi  ,,|'  ,..,ms,.  I„.  i,n.|,.,-st...,.|  tliat  the  i„,,ii. 

■''■'•''  '''"''  '"  ''' '  pi.->Mii-.'  loll..uiii./splaii,.li,ii..  sii,„„lati.,„  is  cans,.,!  1, 

\'''^ siri.-ti.iii    ill    til,-    splaii.-liiii,.    arci    ils,.|f.    as    is   ,>\-i,l,-i„.,.,l    |,y    ,1,. 

''"''    "'■''    ''    ''"'"^  ""'    '"•'•'"■■  "!■  '■'  "Illy    v.M-y    lai„t.   \vl„-ii   th,'  al>,i,>„iiii.i 

''' '  ^•--"■l-<  •ii<'  li'-iiil.-.l  pi-i.ii-  I.,  tl„.  sti„„ilaii..ii  „f  th.'  spla„,-l.,ii,.  ii.Tv. 

Kvii  ;ifl..,-  Iii;ali.,ii  ,.f  ||...  a.|i-..|,al    v.^uis  an. I  .,f  tl„.  1,1, „„1   v,..ss,.|s  .,f  ll, 
splaii.-liiii,-  ai-,-a,  siii„,ilati..„   „f  il,,.  s..!aii.-li,ii,-  iic'v,'   i„ay   sliU  ,-a,is,.  , 

sii'Jil    ,is.-   in   ait.M-ial   1.1 1   pn-ssii,-...   p„ssil,ly  1 aiis,.  s.,"m,.   lilM.i's   „ia 

riiii  IV., Ill  III,,  splaii.-liiii,.  t,,  vasiMilai-  ai.-as  11, it  sitiiat,-,!  uitliiii  tlic  i-.a' - 
of  til.-   spl.-in.-liiii,.   II. TV..      r,,r  ..\aiiipl...   |||.>   |,| 1    \  t'ssi'ls  of  tlw   Iii„il.. 

llll|s,-|.'S. 


t    t 


(11  \|•T 


\  II 


TIIK  CONTIiiM,  dl'  Tin;  f  ||i(  I  I.ATION   iCml'ii) 
CONTROL  OF  THE  VASOMOTOR  CENTER 

Tlic   ,irti\  itiis    111'    III!'    \  i|^ii|lin(iil     fililcT    ;ilc    roMl  I  nl  Iril    pil|ll\     \>y    ImU' 
niniirs  iilicj   pillllx    li\    ,-|lTr|(iit    iiii|iiilscs. 


The  Hormone  Control 

\<  \nlli  till'  li'--pii;it<ii\    fciitc'f,  till'  I'liiif  ImininlK'  is  tlir  li>  i|n>i,'rM  iuii 

•  iin  iitiatimi  111"  tin-  Idimil.  Wlirii  tliis  is  iliricascd,  as  in  iispliN  sia.  tin' 
I  iMPi'iinst  liilii?-  piiit  III'  till'  \  asiiiiiiitiii'  ('('iitiT  lii'i'iiincs  sliinlllatnl,  so 
litiit  llir  IdiHiil  \i"sM'|s  MIC  i-iilistrii'liil  aiiil  tlir  liliiml  pi('-.sUI('  rises.  Tak- 
iii'_',  as  our  ci-ilfriim  nl'  li,\ iltniirii  imi  ciiiiri'iitialiun.  llic  tnisinn  nf  tlu' 
•  .iiiiiin  ijiiixiilc  ill  till'  liliiiiil  I  sn'  pasrr  :i.'i4),  \\i'  iiiav  pmci'i'il  to  invrsti- 
•jiili'  till-  rt'lafiimsliip  liy  olisi'i'viiii;  tlii'  lilooil  in'cssuic  ihiriiiir  I'liaimi's 
ill  till'  I'Milioiiilioxiilc  ti'iisioii  liroiitrlit  almiit  li\  caiisiMy:  tlit'  animal  ti> 
Itiiatlii'  aliiiosplii'rcs  rontainin;;  kiniwii  percent  a  trrs  ol'  the  ann  (Matlii- 
^iiii'  1.  Thus,  it'  a  (leei'ieliiate  efit  is  iiir.ile  to  respire  an  atinosphero 
■ 'iiilaiiiini,'  r>    per   cent    or   more   ol"   earlinii    ilio.xiile.    an    iiiiiiieiliate    rise 

•cTins  in  the  arterial  liloml  pressure.  That  the  inhaleil  eaihoii  ilioxiile 
lilts  li>  raisinjT  the  li\ilroi:eii  imi  eoiieentiatioii  of  the  lilooil  is  imlieateil 
!■>  the  I'ael  that  a  siiiiilar  rise  in  IiIoihI  pressiiie  ean  he  olitaiiieil  li\  iiitra- 
^'  ii'iiis  injeetioii  ol'  a  weak  .solution  ot'  laetie  aeiil  i  "2  e  e.  N  I'li  in  a  de- 
.  l.rate  eat. 

Insteail  ol"  in.jeetinj;  the  laetie  aeiil,  we  may  eause   it   to  lie  pimlueeil 
the  iimseles  of  the  animal   itself  liv    ureatl.v    iliminishin'.'  their  nxxixeii 

'ply       When   a   il lelirate  eat.    fur  example,   is   maile  to   Incalhe   an 

!  iii.isphere  of  almost    iMire   nitrouen,   there   is,  after  a   latent    ]ierio(l   of 
'   lilt  :)0  secoiiils,  a  siiilileii   rise  in  arteri.-il   pressuriv     The  existem-e  of 

•  ^  latent  ))erio(l  in  the  latter  ease,  as  eoiiipaiiil  with  its  .iliseiii'i'  when 
lion  ilioxiile  is  iiispireil,  is  owiiM^  to  the  time  taken  t'oi'  laetie  aiiil  to 
prnilueeil  in  the  iiiuseles  oil   iiei-ount   of  the  oxvLren  ili'iirivatioii.      It 

Mliiirtaiit    to   note   in   the   alio\  e   experiment    that    ileeerelirate   animals 

I'liiployeil   so   as   to   avoiil    the    iieeessity   of   iisini;   anesthesia,    liniler 

'h  the  results  ;ire  tiiiieh  less  detinite.     The  fnet  that  ox>  L'en  ilii>ri\a- 

eaiises  excitation  of  the  vasoeonstrietor  center  has  lieeii   known    fur 

2.17 


I 


I  i 


'£W 


Tin    <rii.  I  I  \Tiiis  ni     nil    iii,im>ii 


s„uu-  liiiM'.  I.Mt  the  r\|.l.ui.iti..ii  tliiit  l.,is  iimimIIv   Ihtm  uivcii  Jiiis  I n  tlur 

it   i»  tllli-  to  II   ilirc.-t   I'lT.-rl   .>r  nxvurll   Hdlll  .III  til.-  .-.litfr. 

Thf  s.nsiiivity  „(  tl,..  , |„|h,,v  ,.,.,„,.,.  •  ,„ar,U  ||i.-  liv.|i'o,t,.,i  i-.i,   , 

iii.iii.v  liiiM.s  uivatrr  lluin  llint  „f  (lir  miIk,  limv  .•.•nlns  in  ||„.  ,,,,i„;, 
••"i.l.  If  nil  iiiiimiil  is  k.pt  tilivr  In  .•iiiiti,.ii,.  ,rH|.ii„li„n  fnr  s,.i,i,.  li,,,. 
iift.T  .Miltiim  tlH-  (•.■ivi.-.il  spinal  rni.!.  tl.,.  su!„i,luiiy  viis..ino|,.r  ...■iiI.m 
will,  as  «..  have  srni,  mailiiallv  ji.-i|iiii..  a  r„uw  a<-ti..ii.  aii<!  tlu-  l..«ri..i 

'•' '  l"''-^'"«'  «  ill  ufiHluallv  lis..  «ua.  i      If.  u  Im,  this  has  I „  atlaiii...| 

th-  aiiiinal  is  ma.l..  to  hivallir  an  atniosphnv  inli  in  .•iirl.oii  ilioxi.l...  a  su.l 

'•''"    '''"''   '"'   ''I '    l'>''^^'ii'«.   will   oiviir,   iMit   to   |.,...lii,-,.   it   a    v..r\    murl 

irr..Ht.T  |Miv,.nta«..  of  this  ifas  nnist  1...  iiis|.ir,.,|  th.-ni  whrii  th..  palli«in 
iM'tH...'!!  Ihr  rhirf  ami  siil.si.lian   ,...nt..|s  is  iiita.i.      \Vlu'r..as  r,  pn-  ....i, 

'•.Hl.on  .lioxi.l..  is  siitlirirnt  to  raiiM.  a  lis.,  of  inrssiir,.  i „„ii„al  h.n 

inu  Its  .-hi.f  Miso,„„ior  ...htrr.  it   tak..s  J.',  p,  ■    ,.,.nt  aii.l  iipuar.l  t ■ 

.III.-.,  a  lil>..  ..n-.Ti  on  a  spinal  animal:  ami  ^  uiilailv.  allhoiiifli  'J  .•.<•.  ,, 
\  I.')  larti..  a.-iil  will  stiiniilati.  tlir  .■lii..f  vasomoi.,  ,rnl..|-,  it  tal«.s  .'.  r  . 
of  N  'J  to  ..\«.il(.  the  spinal conl  (•enters. 


The  Nerve  Control 

llouever  inipoitant   hoiNion..s  may  I.e  in  maintaining  a  toni.-  s^  ..ml;, 

"""    "'■    " •••'••    "'"•    """•<■    ^"iin.'ii    einntres    in    aetivitv    are    mainiv 

'" -'"   =''"""    ^'y   ••'"■'■••'■'"    '"'v    'iiIm's.      The   atTer..nt    imp.ils..s   a,. 

o(    tuo    elasses:      ,1^    (hose    ..ansin!;    a    rise    in    l.loo.l    pressur,.,    eall,.,! 

,,ns,..r.  an.l    .1',   those  .aiisin-  a   fall   in   I.l I   pn'ssur,..  ealh'.!  .h,n;s..„ 

Th..   .llV.t    pr.Ml. I   on    tl...   art..rial    I.I.....I   ,„c.ssure   l.v    stimulation    ..> 

eilh.-r   pn.ssor   ..r   .l.pr.ssor   lil,..rs    is    usually    n...r..   or    l..ss   evaiu-s...,,. 

.•sp..eially  in  ll...  ea '  the  .|,.pn.ssor  fil.ers;  an.l  ^^  I,..,,  t|...  .-han-re  f-.i 

lowinsr  stimulation    ..f  ih..   nerv    pMss,.s  olT.   the   I. n-ssure   al^^a^ 

i-..tiiriis  to  Its  forn..r  i..v..|  This  in,|i,.at..s  that  the  alTerent  ii.ipuls..s  ,'i 
lo.t  air.'et  tl...  toni..  .-onlinl  ulm^h  ll...  vasomotor  .-.'liter  ext'ivisi-s  ..ii  th 
I'loo.l  \..ss..|s.  It  has,  tli.....|oi...  I„...,i  assum...|  l.y  Porter"  that  tli.-r..  a- 
r.'ally  tu..  Uin.ls  of  vasomoi,,,-  ....nters:  ..„..  e.-n.-ern...!  m.-nly  in  ll 
l.riimin;;  alM.ut   .,f  t..mpoi-Mr\    ,,  H.-x  ehan-.-s,  ih..  ..th..r  .-..n.-erii.".|   in  tl 

'"•''""■"•■" 'I    ""•  vas..,ilar  t„,i,.       |t   max    1...  that    the  aetiviti..s  of  ll. 

former  aiv  primarily  ,!-.p..|i.|,.nt   up.>n  a(T,.r..nt   impuls.s.  an.l  the  lati, 

".!•""  '""•"" ^      -lustili.-ation   fo,    this  vi.u    has  h.-en   loun.l  in  ol.s.rx 

tioiis  ma. I..  .„,  t|„.  ,.|r,.,.ts  of  si,,n„|;,,i„,|   ,,f  pivssor  an.l  .|epr,.ssor  lih. 
Ill  animals  un.l.r  the   iiilliieii,  ,■  ,,f  eiirai.    or  al.-oh..l.      With   th,.   form. 

<!rufr.  stimuiaii .f  a   n-.rv..  .....itaiiiin...  ,,   pr,.pon.lerai f  pr..ss..r 

.l.-pressor  til,..rs  pro.liie.s  .|,,i,|,i,.  its  usu.il   etlVet,  l.iit   the  m.-aii   level 

''"'  '•' '  l>""'ssiir..  apart  from  this  ellVet  remains  uiiehanji..,!.     With  ii 

latter   .Ini-    -  .■il,..,h.,l  ..    on    th..    oth.T    haii.l.    th..    retl..x    i-.-sponse   eiilir. 


Tin:    iiiVIHiil     tiK    Till     <  Mil  I  l.xTlnV 


2JW 


.|{<,M|l|H'lll'H,    llllhollttll     it     illlllHMlitltl-ly     l'M|l|l<'llls     U  llt'tl     ill"    lllrillllli     ••fTt'fl 

his  |iiissf<i  o(T,  iiimI  IIiiM'c  is  Mil  I'viili-nrt-  ui'  n  i-li.inui'  in  toii)'  Tlu>  toiiif 
.'iiiii  tlif  ri'tlt'X  iin'cliiiiii«iiis  o''  tlic  vasiiniiitur  rinicr  rnii  mil  llicr'-t'oic  Im' 
I'li'lilirtil. 

At  tin'  |Mi's«'iil  sliiL't'  of  "III'  kiiiiw  li'ili.'!'.  ii  !<•  "Illy  |His>ilili'  fur  us  to 
sillily  th«'  I'ffi'ct  of  sliiiiiihitioii  i.f  |iits,..i-  iiiiil  ili|i»i'Ms.ii  liltcrs  on  tlic 
v.ivdri'tlt'X  cciitiT.  S.ich  lilii'i'N  Jilt'  riihiiiiiicd  in  pnicliciilly  every  sen- 
M,ry  iii-rvf  of  the  liody,  iiiul  it  \»  oiiltj  ii|>|m'iii'  Unit  ii  fiiiil\  iM|iial  inixtiiie 
iiT  liiitli  kiinls  of  lilici-  csisls  in   niosl   of  ttii'se  nerves. 

Pressor  and  Depressor  Impulses,  hepiessor  iiMiulses  ine  alone  present 
ill  tlie  iiirtliih  f/< />;•<. s.Mii'  mm.  Sonieliiios  tis  in  the  liililiit,  lliis  exists 
i>  nil  iinlepeii.leni  nerve  trunk,  oriuiniilintf  liy  two  Iniiiielies.  one  fnuii 
ih.   superior  liirynifeal.  tlie  other  from  tlie  viiifiw,  and  ilese.-inliiijr  elose  to 


I  ■^^  ,- i  I'.ill  „i  I.I.J...I  pi.-.iitr  fiiiiii  .MiLilinii  ..I  till-  .l.-|.ri's-..r  ii.ru-.  Tli.  11111111  u.i- 
'.I  111  ihv  mi. hill-  mI  tlir  tiirvf  ami  tlu*  .Xtitati'ni  iiialiitaiii.'.l  f.ir  t.-v.iili-.-ii  iiiiiiini-^  'I'll,  lin' 
■1.  {.rcHsuff   shiml.l   lie   -III  nun.   I'.w»t  than   hire  *ln.wn.      iFr-.m    lUivli-*-*  ' 

t!  !■  v.iifus  trunk,  to  end  around  tlie  areli  of  the  aorta.  In  other  animals 
''  1  depressor  is  hound  up  with  the  vajius  trunk  from  wliieh  it  can  sonie- 
'iiHi's  he  sejiarated  hy  careful  disseetion.  The  first  piere(|iiisite  in  iiivi's- 
tt-Mlinit  the  cause  of  the  chan<;es  produced  hy  stimulation  of  these  nerves 
i-  'he  elimination  of  any  eliaiice  of  an  alteration  in  heartheat  as  a  result 
1'  simultaneous  stimulation  of  iifTereiit  vairus  liliers.  This  may  he  done 
(  ihcr  hy  PuttiiifT  holli  vairi  or  hv   adniinisteriiif?  atro|)ine. 

Miiiiiilation  of  the  central  end  of  the  cardiac  depressor  nerve  in  smdi 
animal  causes  an  iiiiiiiediate  fall   in  liloud  pressure,  accouipanicd  hy  an 

•nase  in  volume  which  can  he  demoiisl rated  either  in  llie  hind  liliih  or  in 
1    1  of  the  ahdomiiial  viscera-   t'videm f  {.'ciicral  vasodilatation  (  Kijj.  7;t  i. 

'>\  lien  the  central  end  of  a  sensory  nerve,  such  as  the  sciatic,  is  acted 
ii.\  a  stimulus  of  moderate  streiiiitli,  it  will  usually  he  found  that  the 

■   I'ial  hlood  pressure  rises  and  that  the  volume  of  the  linili  tir  of  some 


MICROCOPY    RESOIUTION    T£ST   CHART 

ANSI  ond  ISO  TEST  CHARr  No    2 


_J     ^IPPLIED  INA^GE     Inc 


240 


Tin:  crm  Ti.ATidN-  of  •im;  iii.odit 


IH'- 


.-lliilnlllinill    vis.MIs   iHMnlM.'s   .lilflillisll,.,!       (.vi.lcliro   of  -..ll.Till    VilS()C(.ll.s' 1  i.-- 

Ii"ii.  Hut  vvli.'ii  tlic  sriisoiy  uovM-  is  st iimiliit.Ml  with  cxfiTiiu-lv  wi'iik 
f.iia.li.-    sliucks.    ;ni    entirely    .liir.Tent    lesult    is    likely    to    l.o    olitiiiiie.l : 

""""•'>•  "   '""  "'■  ''1 1  I'l-essiire  mm. I  ;in  ineirnse  in  volume  of  the  liiiili 

'-!•  Niseus.  in.Jie.Mtinir  that   in  this  niiiiuier  we  hiive  stininh.ted  (hi.ressor 
(ilieis.      r.y  e,iivriil   exi)ei-iiii<'iitiitioii   with   <|U;nititiitively   frrji.lnafed  elee 
tii.Ml  stiniiili,  it   hiis  heeii   j-ouiid  hy  Y.iutht  and  others'-  that  on  stiinii- 

hitiim-  Ml  atr..reMt   nerve  with  weak  shoeks,  a  fall   in  hi 1  pressure  is 

the   lirst    elVeet    to   I l.served,   and   that    this   he.-oiiies   re  and    more 

niaiked  as  the  strength  of  the  stiniidi  is  inereased,  until  a  eertain  ojiti- 
mum  is  reaehed.  after  whi.d.  the  fall  in  l.h.od  pressure  heeomes  less  evi- 
dent. \Vli..ii  a  eert-iin  stren-th  of  stimulation  is  exe.^eiled.  a  rise  instea.l 
"'■  '1  '■.dl  .M-.Mtrs.     After  this  p.mit  ad.litional  inerease  in  stimulation  eauses 

■'■  ••""!   """•'■  iiiarkci  .■levation  of  hh.od   pressure  throu-h   a   verv   Ion- 

ran^ri'  of  stimuli. 

Stimulation  of  tw,,  atVerent  nerves  at  the  same  time  usually  produces 
a  -rreater  rellex  vasoim.tor  ehan-re  than  the  stiiinilation  with  an  e<|uiva- 
li'iit  stren-th  of  eurient  of  either  nerve  alone.  That  is  to  say,  the  eff.'.-t 
in.Hlueed  hy  st imnlat in-  the  eentral  en.l  of  hoth  seiaties  simultaneouslv 
udl  he  -reat.'r  than  that  pro,lueed  hy  stimulating  either  alone  with  douhlV 
the  stien-th  of  stimulii.s. 

•^^  ''''^  ' "  •^'='''■'1  id>ove,  the  reflex  elmnsre  in  l.lood  pressure  is  often 

Muite  transitory  in  nature,  althoufrh  the  stimulation  of  the  pressor  nerve  is 
liiMiniained.  When  this  <leeline  has  oeeurred.  the  pressor  reaetion  ean 
often  he  renewed  hy  shiftiim  the  stimulation  to  a  seeond  nerve.  Tiiese 
fa<'ls  eonrernin-  the  greater  eftieaey  of  eond.ined  stimulation  of  several 
nerves  are  of  eonsiderahle   importance   in   eonneetion   witli   the   (.'eneral 

ipiestion  of  relies  elian-es  in  hi 1  pressure.     For  instance,  many  of  the 

IM-essor  tihers  found  in  the  sciatic  nerve  are  connected  with  the  recei)tors 
that  mediate  the  sensations  of  the  skin.  When  tliese  receptors  are 
stimulate.!,  as  l.y  heat  .,r  c.l.I,  rellex  ehansjes  in  hlood  pressure  occur 
(pr.>ssor  reacti..nt.  i  i'ii:.  74i.  and  it  is  inip..rtant  to  rememher  that 
lo.'.'dize.l  stimulati..n  ..f  th.'  skin  is  less  etH.-ient  in  hrin-jiufr  fd.out  su.di 
vascular  chan-vs  than  stimidathm  api)lied  over  larjic  areas,  even  when 
Ih.'  I.ical  stimulus  is  intens,'  an.l  the  -eiieral  stimulus  mihl  in  character. 
.Fui.ipin-r  int.,  a  m,..leral,'Iy  c.i.l  hath  will  cause  a  much  j^reater  rise  in 

•■"■''■'■i"'  ''1 1  pr.'ssure  than  plu.i-in.j'  the  hnn.l  into  ice  cold  water. 

Mechanism  of  Action  of  Pressor  and  Depressor  Impulses.— When  we 
•  •onsi.ler  the  exa.'t  mchanism  hy  whi.'h  thes.>  alTcrent  impulses  o]).'rat.-. 
we  liave  t.)  hear  in  min.l  f.,ur  po.ssihilities:  the  rellex  fall  produce.l  hy 
stiinulati.i!)  of  a  .h-pn^^-v.,.,-  afTereiif  fiher  may  he  due  either  to  a  slimula- 
ti.Mi  ,<{  the  va.s.,.lilator  part  of  the  center  or  U>  an  inhihiti.m  ..f  the  t.,n.' 


TIIK    CONTROt,    OP    TlIK    CIRCrl.ATION 


241 


(,l  the  vasoconstrit'tin'  p.-irt;  nml,  ('oiivpi'scly,  a  rise  in  arterial  pressure 
caused  by  vascxMiiistrietiou  may  lie  dependent  eitlier  on  a  stiinulatiim  of 
flie  vasoeoiistrietor  part  of  tlie  eeuter  or  on  an  inhibition  of  llic  tone  of 
the  vasodilator  part.  All  of  tliese  chanses  have,  as  a  matter  of  fact,  lieen 
shown  to  occur,  at  least  under  certain  conditions,  althouffh  the  evicfence 


I'i«.  74. — The  rITcct  of  stritHK  stiiniil.-itiiiri  (hi-.il)  ot"  Ihc  >kin  nf  tin*  fn.)t  on  lln-  .irttTi,iI  ltlu<i.l 
irt^^iire  and  rt  spii  atnry  iiHtvtiiunts.  I'l'juT  triuing,  thoracic  tnovt  iiu  iil ;  I'>w<t  traciUK,  artiii.il 
lit. .11,1    iirfssiirc. 


lor  the  iuhihitioii  of  dilator  tone  is  as  yet  a  little  uncertain  (see  Fi}r.  71)). 
Without  goinf;  into  the  sultject  in  detail,  we  may  nevei'theless  take 
its  an  example  of  the  iiiel'iods  liy  wliich  the  infoi'iiiation  lias  lieen  oli- 
taiiu'd,  the  experiTnent  perforiiied  Iiy  IJayliss,'^  showing:  that  the  ^asodi- 
l.itioii  which  results  from  stiiiuilalioti  of  the  depressor  iicrve  is  owitii; 
partly   to   removal   of   vasoconstrictor   tone   and   pai'tly   to   vasodilator 


It       i': 


i 


'2V2 


Till    ciHcriATiov  DV  Tin:  ni.non 


stiniuliitidii.  Tlic  vdlnnii'  df  the  hind  liiiil)  ..f  a  curaiizfil  .•uul  vaKotoiuized 
rabl.it  iiicreasos  \»li,>ii  tlic  coiitial  oiul  of  tlio  canliae  ilciirrssor  nerve  is 
stimulated.  In  order  to  dctfiinine  whether  tliis  dilatation  is  due  sololv 
to  the  removal  of  vasoeoiistrietor  tone,  tlip  aliove  ex])erimeiit  was  repeated 
on  ar.diliif  in  ^^hieh  the  s\  mpathetie  chain  had  lieeii  eut  below  the  level 
of  the  seeond  lund)ar  spinal  roots.  By  sueli  an  operation  all  the  vaso- 
eonstiietor  fillers  to  tli.'  vessels  of  the  hind  limb  are  severed,  but  the 
vasodilator  libers,  sinee  iliey  emanate  throuirh  the  sacral  .sensory  roots. 
are  left  intact.  It  was  nevertheless  found  on  stimulatitii^  the  depressor 
nerve   that    dilatatinn   of   the   jiind    limb   still   urred,   thus   indicatiiif; 


I'ijkT.i'n    .|i..>M!,K    III,'    |ii,,l,;,l,l,.    .iir.iiik-,  ir,  ul: 


•  ilMr    riHixi- 


HSU, 


■i    M.1-.  If   M   ,111,  ri. 4,-. 

/'.    \.l<',.lil;,i.,r   II,  ,v,-   i;l„r   t,rr,;ii:,lniit   „ii   .-(   a:i,l    inlul.ilini;   it*    iMlural 


t'lmis,    as    illilicat,  li    liy    - 
tor   arise 


iru.n    111,-    ,l,i.i,,,r    ,,-,.,r    i.hi    ami    th,     ,,,nstr„i,,r    ,,nl,r     (ft)     r.spcclivdv. 

I-.   AtUvrvt    .|,|.r,—  r    M-.i,    .i,M  hni;    ,:;„,    ,„,,    !,,„h,,.    ,„„.    „,     „l,ia,  '(-)     i„hil,it,    th,-    r„  , 

i^7:u::T;.u!:L''"  '■""' ' '  '^""- "-  ■'''■" -  -usi„« .iiiauii,,;  of ;;:;  aii;':i,:  a':, 

of  l^i:„'i",^^,n';.''  """"-    '''-'  ^"■'   '^''"'"'-«  ''"'■   '"■>   •■.■., fi.r,.  ca,isi„«  vas.„-.,ns,ri.t,nn  ami   r,s,. 

.i.,":.,!r\?ri;'.i';.;::'r  ""■  '"'""^  '^  "■•  •'-'  -"  ''•■■—-  •— -  -"■  "■•■  •«-.„, 

that  stimulation  throu!.'h  vasodilator  (ibers  must  liave  taken  place.  Con- 
versely, in  aiiolher  experiiiient.  i-isfead  of  the  synipnthotic  chain,  tiie 
spinal  cord  was  cut  below  the  level  of  the  second  lumbar  segmort.'thus 


I 


<135 


Tin;  roNTHMi.  oi'  Tin:  ciik  ir.vTinv 


243 


si'VcriiifT  tlip  tliliilni'  Imi  not  tlic  ('(Piistrictor  p.itli,  miliI  jiijiiiii  (Icurcsscn- 
slinnilatioii  ciiiiscd  tlic  vuliiiiu'  ni'  tlio  liiiili  Ici  iin'ii'iiNC,  iiulii-atiiij;  llial 
an  inhibition  of  cunstiii'lor  toiu-  nnist  liuvc  occuricd. 


I  -4 

i     I 


3 


■a 


Reciprocal  Innervation  of  Vascular  Areas 

If  ninst  not  lie  iniasiincd  tliat  (•]ianvr<'><  in  tlie  ralilicr  ot"  tiio  lilootl  vcs- 
>.cls  occiM'rin!,'  in  one  vascular  area  aic  ncrcssarily  occui-iiiij;  all  o\  cf 
the  liody.  On  tlu'  contiaiy.  a  most  inipoitant  icciprocal  felationship 
exists  in  till'  lilooil  sMintly  to  dilVcrciif  parts.  Afto-  fond  is  takai,  ior 
example,  more  Mood  is  i(i|iiiicd  by  the  diLresti\t'  oi'irans  than  when  they 
ai'e  at  I'est.  and  this  is  insured  liy  dilatation  of  their  own  v<'ssels  aloiii.' 
with  t'ceipioeal  eonstiii-tioii  of  those  (d'  other  par's  of  the  bod.v.  (*n 
aeeount  (d'  'he  relatively  yreat  eapaeity  of  the  nhilominnl  xcsse's,  their 
dilatation  <.urin>;'  diu'eslive  a<'ti\it>  is  usuall.\  u'reater  thaTi  the  reeiproeal 
eimstrietion  of  the  otluT  vessels,  so  that  the  diastolic  hlood  pressure  falls, 
iiccessitatinir  a  more  powerful  cardial"  dischaiire  in  oi'dei'  to  maintain 
til"  meai;  jiressure.  After  taUiiii;-  I'ood,  the  systolic  picssme  does  not 
as  a  rule  fall  so  much  as  the  diastolic,  if  it  falls  at  all;  and  the  ]>ics- 
sure  ]>iilse  theret'ore  lieconies  jrieater  and  causes  a  <,'reater  live  load  to 
he  applied  to  the  vessels  with  each  heartlieat,  Dlirinu'  the  sudden  strain 
that  is  tlirown  on  them,  wealxcncd  arteries  may  tri\e  way.  especially  in 
the  hralu. 

Anotltei"  examiile  of  reciprocal  action  of  the  vascular  system  is  seen 
ill  nnisciilar  r.rciri.si.  The  vessels  of  the  ;ictive  nuiscles  dilate,  wliile 
tliipse  elsewhei'e  eonsirict.  The  local  dilatation  in  this  case  is.  however, 
not  entirely  ;it  least  a  nervous  )>heiioiiieniin,  heiiij;  ejiused  in  fact,  as  we 
shall  see.  Iiy  horiiioiie  .ictioii  on  accniiiit  of  the  ioc;d  increase  in  hydro- 
i;vi|.ioii  concentration  i  sec  p.iyc  414 1.  Theiv  can  he  little  doiilit  that 
loriiJ  irritatifft  to  the  surface  of  the  liody,  such  as  hot  applications,  lini- 
iiieiits,  etc..  act  in  the  same  wa.v;  they  cause  local  dilatation  of  the  su))er- 
licial  and  jierlijiiis  of  the  immedijitely  underlying'  vessels  and  eonsti'ic- 
'ion  of  those  elsewhei'  in  the  liod.v .  A|)])lica1ion  of  eold  to  local  areas 
oi  skin  similarly  causes  local  const rictioii  accomi)anio(l  liy  reciprocal 
dilatation  elsewhere.  This  action  of  cold  is  very  marked  in  some  parts  of 
llie  liody.  such  as  the  hands,  ulierc  li\  Sleunit's  method  I  pa  ye  'JS:',  i  it 
eaii  bo  shown,  not  only  that  the  bloodflow  of  the  hand  to  which  the  cold 
is  applied  is  fil-eatly  curtailed,  hut  also  that  of  the  opposite  .side. 

Experimental  denmiisti'ation  of  reciprocal  vascular  innervation  is  fur- 
nished by  numerous  experiments.     !('  the  centi'al  end  of  the  v'reat  aui'ic 


uiar  tiervu  o 


f  11 


le  c;ir  is  si  iiuuia.ed   ni  a   rati 


bit. 


1   hi;inctiin;r  ot    itie   vcs 


f  thi 


i   n 


sels  of  the  ear  oeeurs  ;it  the  same  time  as  a  ri'C  in  arterial  blood  pre? 


Ml 


Tiir:  ( ri{(i  i.ATiox  uv  Tin:  iilood 


sine  (I.c,\.'.|i  ivllcxi.     Siiiiiliiily  w  lien  tlic  iTiit rjil  cn.l  of  ..ii<>  of  U)p  sen 
"ly   inc. Is  i>(  till'  Iry  of  ii  iloy;  is  st iiiiiiljit..,|.  tliciv  is  a   lis."  in  aitfi-iiil 
Modd  jni'ssiiiv  ami  an  incicasc  in  tin'  volmnc  of  the  iini!). 


THE  INFLUENCE  OP  GRAVITY  ON  THE  CIRCULATION 

If  til.'  ait. -rial  Mod. I  prcssiiro  is  nicasiiicl  in  the  arm  ami  Icl'  in  a  man 
slan, litis;  .'ic't,  a  .lilV.'ivm-.'  .•oiT.sp.m.Iiinr  t..  tlic  liy.ln.static  .'rtVct  of 
gravity  will  Im'  loiiml  l-i'twcn  tli.  tw.i  i.'a.Ilniis,  In  .■oiiipaiison  with 
111.'   hi-h   pnssnic   mniiially   .'xistini,'  in   tlic   ait.'ii.'s,   tliis  .linVivn.'.-    is. 

liow.'v.T.  of  littl.'  siifiiili.'ai On  til.-  ..tli.T  liaml.  in   tin-  vins.  wli.'iv 

III.'  a\.Tayv  pn'ssiir..  is  low.  -lavity  woiii.I  calls,'  s.'iioiis  ciiil.airassiiicnt 
to  till'  cil-ciilatii.ii   of  l.loo.l   were   it    not    for   llic   valves  and   tlu-    f..ic.'s 

\\\\:i'h  iiioM-  III.'  1,1 1  JM.yoml  til, 'Ml   (pai,'."  1214). 

Ill  ,'iv<'l  aiiiiiiais  til,'  part  ,.f  tli.'  ciiviilation  in  wliicli  l,I,„.,l  miirlii  sta- 
iml.'  .'IS  a  r.siiit  of  u'lavily  is  tlic  splam-Iinic  area.  Were  sueli  sta«rii;i 
lion   t,,  o,-,iir,  til,'  MihmI    w,Hil,l   not    I,.-   r.'tiirn.'.I   to  tli.'   rij;Iit    Ii.'art.   so 

lliat   til.'  art,'ri,s  \\,,ul.l   not   r.'.-.'iv.'  siifli.-iciit   I.I |  t,i  maintain  an  a<lc- 

,|ii.'it.'  .•ii'.-iilati,,n.  iparti.'iilarly  in  tlie  vcss.'Is  of  llie  Iiraiii. 

Si„i|,l.'  .■\p,'ri:ii.'iits  .Icviscl  I,y   l.i-onar.l   Mill''  -    illustrate  these  priii- 

'■il'lcs.      Wlh'ii   a   siiaN,'.   for  .'xaiiipl,-.   is  pinm'.l   ,,nt   on  a   l..n^'  ])i,., f 

" '  ■■I'"'  •■I"   "I'l'iiin-r  Ilia,!'    opp,,sit,'   tli.-  Ii.'art.  tliis  ..riraii   can   I.e  sc.'ii 

to  lill  a,l,',pial,'ly  with  I.Io,m|  as  loii-  as  tin-  animal  is  maintain.',!  in  the 
hoii/,.iilal  i)osili,,n.  When  plae.'.i  vertically,  however,  tli,'  heart  l>c 
cmi.s  l,l,M„l|,.ss.  If  now  th.'  tail  .'ml  ,,r  th,'  animal  is  pla.-.'.l  in  a  ,'yliii,|.'r 
of  ual.r  s,,  ,is  t,,  ov,'r,-,,mc  tlu'  ,'ir,',-t  of  -ravity,  the  heart  will  he  s.'.'ii 
I.I  lill  .luaiii  uilh  IiI.mmI.  l-;vi,l,'iitly  in  sm-h  all  animal  ther.'  is  iiii  iiie.-han- 
isiii  111  .•,imp,'iisalc  f,)i'  yra\it.v. 

If  a   ,Iom.'sti,'   rahhit   with  a   Iar-_'c  p,'n,liiloiis  ali.|om.>n   is  li.'hl   in   th.- 

v.'i-li.-al    lail-,l,.un   posiii.,ii.   sta<riiatioii   of   hi i    i„    -lie   splanchnic   ves 

si'ls  o.'.'iiis  to  sii.-h  an  ext.'iit  that  in  from  lifi.'.'n  to  tw.'iity  miniit.'s  th,' 
.■iiiimal  .11, 's  fr,,iii  .'.rel.ral  anemia.  If  an  ahihimiiial  himler  is  (list  of  all 
appli,',l.  th,'  v.'rti.'al  ]i,.siti..ii  will  not  hav,'  tli.'  sain.-  coii.s.'.|ii..n.-.'s.  This 
cxp.'rim.'iit    illiistral.'s  .-l.'arly  the  possihle  e\  il  .'ITccls  that    -ravity   may 

pi'o.lii.-e  in  animals  in   «  lii.-li  n.>  m,',-liaiiis xists  I .iiiiiensalc  "f,.r  it. 

I'la.iii!;  an  animal  such  as  a  .l,,-.'  umh'r  liirht  .tli.'r  anesthesia  in  tlic 
v,'iti<'al  tail  ,l,.\\n  jiositioii  i.ro,Iii.'.'s  an  iiMm,',Iiat,'  fall  in  arterial  l.l.,.,i| 
pivsMli'.',  as  shoun   in   the   tra.-iii!.'   i  Kiu'.    7(ii,    follow.'.l    l,y   a   certain   ,I,'- 

-'''•'■  "I'  ••<iiiip,'iisati v,'n  whil,'  th,"  animal  is  still  in  the  erect  position. 

The  .'xli'iit  t  '  which  this  .■oiiip,'iisati,>ii  o.-.-iirs  vari.'s  with  the  .lepth  of 

the  anesth.'sia.     If  i! xperim.'nt  is  r.'pcat..,!  after  a.lmii.isicntiir  a  iar<r.' 

.lose  .,f  .•hlorotorm.   n,it   ..iily   \\\l\   the  initial    fall  I.e  iiiiich   fircater    I.iit 


9* 


nil:  roNTiioi.  riK  Tin;  (iKiri.ATniN 


liJ") 


>illisr(|lli'nt    rci|ll|>i'risiitii>ll    will    lie   |i|-!irt icilllv  illisrlit.      'I'lic  ;i|)|ilii-;iti'iii   of 
llii'sc  I'licfs  in  the  npcijiliiin'  iihuii  will   Im-  M-H'-cvidi'iil. 

I naid   Mill   li;is  sIkiwh   Hint    llin'c   rnct'irs  iiic   iiiMiivcil   in   the  coni- 

in'Msiitillj;   liiccliiiliisiii:    (1)    tlio   tiiiiicity   of   thi'   iiliilKiiiiniil    iiiiis;iil;iturc ; 


ill 


//^::/^' 


,Vvv'  "■'•^'■^''^''•"■^^"•'•WvA'i' 


'  ■■■    ""•      A. .nil-    I.I i    |.r(«-iir;-.    '.Ii.,»im:    III,-    ,(T,,i     ,.f    |...vti!r.-:       .f.    virli..,!,    Ii.a.l  Ml.;    /(      lioii 

/'■rilal:     I.    virli.al.    In  ,i.l  .i..wii ;    /',    h.,i  1/..11I.1I.       I  [..II. I 

_''  till'  tone  of  tlif  s|)l;mrliiiii'  lil 1  \,    -i-ls;  Cii  ihi-  ]niiMi)iMir  action  of 

ilif  ivs|iiriilory  inovciiic'iiis.  The  iiii|ioit.iiii-f  of  the  lirstiin'iitioiicd  t'.-n-- 
t'lr  can  Im'  icadily  show  ii  liy  iiiakiii";  a  crucial  incision  of  llic  aluloin- 
liial    walls    in   an   animal    in    tiic   erect    position    (  {'"i;;-.    77  >,    and    that    of 


I-1W      ;:       Ti.Mim;    1,,    ,!„,v,     ihr    ,tr,-,t    ..f    i..,:nil>     ,,„    III,     ait,  ,i.,I    I, I |     ,,,..,,,  \,      (      ,|„ 

''    "■'-    I'''""'    '"    ll"-    v.rlu.il    |...,li..ii;    .,t    II.    ii„     ;,1,.',,„„„    „  ,,    ,.,„Mr,.-,.|      .,i     i        ,    ,iu,,il 

■'    r,,,.|    t,,    ||„.    |„,ri/.,iit,il    ,,.s,t|,,„.       i|.-,,..„    I,,.,,,,,,,   I     11, |]   , 

•lie  siM-i.nd  factor  hy  eiittim,'  tin-  u'lt'at  s|.lanclinie  nerves.  „y  ilie  spinal 
"111.  After  such  an  operation,  even  uliile  in  the  aoii/mital  posititm.  as 
'■  liave  si'i'ii.  the  lilood  pressure  falls  to  a  eonsider.aMe  estcnt.  If  the 
liiiiial  is  now   jilaced  in  the  vertical  tail-down  ixisition.  however,  it    falls 


'It, J  J 
if  • 


1 11 

..  1 

■'1- 

!■ 

i' 

'  \ .' 'li 


f 


:\, 


i, 


hi  m 


p  iiii  I 


i-i  t 

4  \ 


!  ! 


11. 


w 


i  m 


!i      i 


24(i 


THK  cnm. ATioN-  or  tiii:  iii.doD 


to  the  /cri)  liiii-  iiml  tlic  .■iiiiiinil  scmM  difs  ( I''i'_'.  "Si.  'I'lif  iiilliiciicc  of  tlic 
(liinl  I'iii'tor  i>  not  so  urfciit  ns  of  the  other  two.  Imt  i-jiii  lie  shown  li.\-  thr 
incit'iiscd  rcs|iiiatoi  \    iH-tixifv  wliich   is  likely   to  (!e\clop  in  the  vei'tieiil 


If     rs.     The   cffiit   nf  uravity   on  tlie   aortic   pressure   after   ilivNinii   nf   tlie   s|>irial   conl   in    th. 

;ii'|..  1    .lui-.i!    rtKiciii.       I'.y    iil.uiiij:    llip  .iiiTiii.ll    ill    the    veniial    feet  ihiwii    |ii.-tiile.    the    |ire«»iire    fell 

.ilni.i^l   1,,  /,  1,1,   l.iit   nil   reluiTiiiii!   It    |.i  the   hori/..iit:il   |...-liire,   the  lirtnhiti.Mi    «;i>   resliire.l.      (Fiopii 
1.1  "11,11,1    llill.l 

tiiil-douii  position,  the  iinciuic  eoinlition  of  the  i-esiiii-atoiy  eenter  lieiiij: 
no  doubt  tlie  ciiiise  of  the  inereiised  ii'sj)iiatioii. 


[4| 


CHAI'TKI}  XXVIII 

I'KCUIJARITIKS  (»!•'  I'.I,<><»I)  SFI'I'LY   IN  CKUTAIN  VISCKRA 

I'p  to  the  present  we  liii\t'  liccii  (•(iilsidci-iiis,'  tlic  ciicilljltinii  of  tho  I)1<>"<1 
tVdin  a  Boiioral  jioiiif  »{  view,  TIiimt  nro  ccrtiiin  Di'jriins  .-iiid  tissuos.  liow- 
rvcr,  ill  wliicli  tlir  L't'nci'iil  inccliaiiisni  is  nltort'<l  in  ordi-r  to  iiippt  \tc<'\\- 
liiir  io(|nircnicnts  (if  Mood  snjiply.  Tims,  it  is  evident  tliat  the  l)iniii, 
iiieasod  as  it  is  in  tlie  riyid  eriiniiiin,  will  Ite  uiialile  to  eontract  and 
expand  as  a  result  oi'  vasoeoiislrietion  or  vasodilation.  On  the  other 
liaiid,  we  know  that  the  iilood  supply  to  this  (Ufjaii  does  vary  ooii- 
sideraidy  from  timr  to  tiine.  What  is  the  nature  of  the  ineehanism  Ity 
"hieh  siudi  ehanjies  aie  hron<.'ht  aliout  ?  In  the  ease  of  the  liver  tiie  eir- 
iidatiim  is  i)e<-uliar  on  aeeount  of  the  fact  that  Mood  is  earried  to  the 
or-ian  by  two  vessels,  in  one  of  "hi<-h  it  is  supplied  under  hiffh  pressure 
iiiid  in  the  other,  under  lo\v  ])ressure.  AVe  must  investiKate  the  rela- 
tionship of  these  two  sources  of  liloiid  supply.  The  eireulatiou  through 
the  coronary  and  luilmonary  vessels  must  likewise  receive  sjieeial  atten- 
tion on  aee<)unt  of  the  hiirhly  spei-ializeil  functions  of  these  organs. 

THE  CIRCULATION  IN  THE  BRAIN 

Anatomic  Peculiarities 

Serious  eurtailmeiit  of  the  lilmxl  .«»/)/</(/  to  the  hrain  is  v'uarded  ajrairist 
liy  the  existence  of  the  eii'cle  of  'WiHis.  besides  the  foiu'  main  arteries-- 
tlip  vertehrals  and  the  two  carotids-  the  spinal  arteries  contrihute  to 
tiie  Iilood  supply  of  the  circle,  and  eonseipiently  in  certain  animals,  such 
IS  the  doir,  the  four  main  arteries  may  lie  lijrated  without  causing  death. 
In  man,  however,  ligation  of  both  carotids  is  usually  fatal.  The  free 
iiiastomosis  displayed  in  the  circle  of  Willis  is  not  maintained  in  the 
I  iise  of  the  arteries  whi(di  run  from  it  tn  supply  the  hrain  structure.  On 
Mie  contrary,  these  vessels  are  more  or  less  terminal  in  <'haractcr;  that 
is  to  say,  the  capillary  .system  pi-oduceil  liy  the  ditTerent  vessels  does  not 
I'reely  anastomose,  so  that  the  olist ruction  of  one  vessel,  or  an  important 
iiraneh,  is  followed  Iiy  death  of  the  suiiplied  area.  The  vesse's  which  go 
to  the  pia  mater,  however,  lireak  up  into  numerous  smaller  branches, 
which  freely  anastomose  before  entering  the  brain  tissue. 

247 


i 


fm 


LM8 


TIIK   IIHCIt.ATION    ()!■'   TIM:    m^KHl 


ir    1 


Tho  v.'n.MiN  Mooil  is  n.l'.rtr.l  I..V  tlic  small,  v.-ry  fliiiiWHllfd  aii<I  valvo 
Ifss  fcirlii-iil  veins.  These  rmi  together  to  form  liiijjer  veins  dis- 
ehiirciiiu'  into  the  sinuses,  the  o|ienintis  into  whieh  me  kept  i>ateiit  l>v 
tlie  aiiariiremeiit  of  iluia  mater  aroiiml  tiie  orifices.  The  sinnses  exist 
lietween  the  diira  and  sluilj  and  are  so  eonstriieted  tliaf  tliey  can  nol 
lie  c(>m|)res.sed.  parlicidarly  those  at  the  Itasc  of  the  hrain.  l-'rom  th.ni 
tlie  Mood  is  eonve.ved  niaiidy  to  the  internal  .jntMilar  vein,  some  of  it 
Iniwever  cseapintf  hy  the  anastomo.ses  cxistintf  hetwren  the  cavernotis 
sinus  ami  the  opththalmie  veins,  and  Iiy  the  venous  i)lexus  of  the  s])inal 
cord.  The  most  striking  peculiarities  of  the  veins  are  their  patulous  con- 
dition and   the  :ili>en. f  valves,  s,.  that  any  chantje  in  the  IiIckmI   \mv 

sure  in  the  iiiternid  .jntiular  vein  nnist  he  immediately  ri'li.-cted  in  that  of 
I'l"'    ^''' >*   sinuses.      This   explains    why   compression    of   the    abdomen 


i;;::;;■:!;:v^"'■;l;:v::''::;;;^^^;':'HJ:• ;"::;.;;  J::^  '" 

I  lr..ni   II,,,,, M\  /';>„„/,.,,,,  I  I      J  11      I 


l.l,....l 


I'l   ml,,   till 


causes  venous  iilood  lo  i|ow  from  an  openinf;  made  in  the  longitudinal 
sinus. 

•' nsidcrinj;  the  cereliral   circulation,  ;inolher   factor  that    must   he 

lioine  in  mind  is  the  iireM'iice  of  c,  n  hrospliiiil  /hilil.  This  is  contained 
in  the  suharachiioid  sjiaces  of  the  l.iain  and  spinal  cord,  these  sjiaces,  in 
the  ease   of   the   I. rain,   heinj,'  often   consideraMy   enlarf;e<l   to    form   the 

eistcrna-.     Tl crel»rosj)inal  llui<l  is  also  present  in  the  ventricles  of  tin' 

hrain,  which  it  will  l.e  rcmemhercd  communicate  with  the  .siiliaraehuoid 
.si>aces  tlirou«:h  tin-  foramen  of  .Masreiidic.  etc.  It  is  unlikely  that  the 
eeiel.rospinal  tliiid  is  of  much  importance  in  connection  with  the  control 
of  the  hlood  supply  to  liic  l.rain  ti.ssue.  It  may  ho  nu'iely  a  luhricatiii}; 
fluid:  at  least  it  is  so  small  in  amount  {CO  to  W)  c.e.  in  man)  as  to  1.^ 
apparently  of  little  value  in  hrin-.'inj,'  about  an  alteration  in  brain  volume. 


■■iicn.iAKiTir.s  OK  iii.iNiii 


■II. V    IN    I  IIITAIN    Visi  IU\ 


M9 


Althoii^li  iioriiiallx  >»i  Kcantv,  its  scci'i-timi  fiiii  Ih'chmic  n'liiiirkiililx'  slim- 
iiliitcil  iimlcr  crftiiin  (MiiiililiiiiiH  jis  in  I'liirtiirrs  of  tiic  linsc  iif  tlic  sl<ull. 

rmlfi'  tlics iiKlitiiiiiH  ill  iiiaii,  it  iiiiiy  diaiii  away  at  the  ratu  of  about 

-IiHi  c.c  a  (lay  or  nmro. 

Till-  lliiid  is  appai'i-iitiy  sri-n'tctl  fr  mi  tin-  clioi'iiitl  )iifXUH,  fur  wlirii 
ilic  j)atli«avs  l-y  whidi  tlic  Nciiliii'lcs  roniniuMicatc  with  fiu'  siiliararh 
Muiil  spncc  arc  olistiiicti'il.  it  collects  in  tlie  vciilrieles,  ])ro<|iieiii|;  internal 
livilroeepliallis.  I'mler  certain  eoiiilitions  its  alisorption  is  also  very 
vapiil,  as  show  II  e\pei'iiiientally  li>'  the  rapiditx  with  which  ph.\  sioloiiic 
^.iline  is  ahsorhe(l  when  it   is  injeeleij  into  the  snhainehnoiij  space.      This 

;ilpsoiptioii   is  heliexed   to  ur  thlouudi    the   I'Mcihioiii.-in    liiiilies,   wliieh 

.'ii'e  minute  saelike  iirotrusioiis  of  the  arachnoid  into  the  interior  of  ,1 
MMoiis  sinus.  The  ineinlirane  that  separates  lilmid  and  eerelirospinal 
itiiiil   is  extreme. y   thin  at    these   ])lai'es   1  l'"iy;.   TM', 


Physical  Conditions  of  Circulation 

(•n  aeeoiiiit  of  these  anatomie  peculiarities,  the  jihysii-al  factors  con- 
iinilin);  tlio  cirt  uliitioii  of  Mood  to  the  lirain  are  considerahly  dMTerent 
tiMiii  those  ol)tainin)r  in  any  otlier  part  of  the  liody,  with  the  jiossilile 
i-M-eption  of  the  liones.  In  other  vascular  areas,  we  have  seen  that,  when 
ililatation  or  eonslrictioii  of  the  vessels  occurs,  a  marked  increase  or 
iliiiiinutioii  of  the  volume  of  the  part  lieecum's  evident,  Siicji  a  chanire 
in  volume  is  evidently  impossilile  in  the  case  of  the  inain  hecause  >( 
'lie  rijiid  <'raiiiu-i  in  which  it  is  contained.  In  fact,  from  a  physical 
l"iinf  of  view  we  must  consider  t!ie  lileod  ncsscIs  of  the  hrain  as  ]>ro- 
.itctiiiK   into   a    rijjid    ease   tilled    with    incompressililo   material.      I'nder 

Micse  eoiiditions  it   is  oltvioiis  tliat   the  \cssels  as  a   wliol uld   licit  her 

ii'iitract  nor  dilate  without  some  increase  oi'  decrease  in  the  volume  of 
Mie  contents  of  the  cranial   i-avity   (Leonard   llill'''K 

Snnii'  have  thouj.'ht  that   tiie  eerelirospinal  Illiid  as  it   llows  into  or  out 

■r  the  spinal  ecu'd  miiriit  accomplish  tliis  alteration   in   the  cranial   con- 

'   Ills,  hut   tlio  relatively   small  amo\iiit    of  availalile  ccrehrospinal   tliiiil. 

'lie  smalliiess  of  the  oix'iiiufrs  liciucen  the  Inain  and  the  spinal  ecn-ij,  and 

'iic  lack  of  experimental  eviilence  that  such  i-hanires  in  volume  of  cere- 

I'liispinal  lluid  in  the  spinal  cord  do  actually  occur,  all  stand  in  coiitra- 

iiclion  to  such  a  view.     I[o\\c\i'r,  although  the  vessels  as  a  whole  mii,'ht 

'  "1   eoiitraet  or  expand.  .\ft  some  vessels,   like  the  jirtcrie-i.  mi>rlit   con- 

'   act   simultaneously   with   a   correspoiKliiis,'   dil.itjition    of   other    vessels. 

ich  as  the  smaller  cerehral  veins.     In  adniittin<!;  the  possiliility  of  si  iiu 

'  ciprocal   relatioiislii|i  hetwcen   arteries  and   veins,   we  must    remem  icr 

•!.it    it    is   oni.v    hefore    the    wcll-)irotected    sinuses    are    reaclieil    that    a 

'  'i;inge  in  the  ealiher  of  the  veins  would  lie  possible.     Hut  it  is  dinicult 


I 


\i     if 


!.J0 


Till     r  IHi  I  I  ATliiV   OK   Till:    lll4Mit> 


>i 


V 


III   si'i'   liiiw    slH'li    rrripi ill    ililiililtinti    iiiiil    (•iiiistrictiiill    could    he   of  (lliv 

iiihiiiild'jc  (Ai-fiii  |H'ili.i|(s  ill  Diiisiiiif  I'crtiiiii  iin-iis  to  rcpi'ivc  nuni 
Mood  tliiiM  olliiTs  A  iii'i|)iociil  irlntion>liip  iiii|;)it  niso  cxiNf  hctwoin 
iiiljiMMiit  iiitii  ii.l.'s  lis  Mel!  IIS  liilwfcii  jirtcrlolfs  niul  vi-iiis;  wlicii,  for 
cMmipI"',  till-  iii'Mi  ('('liter  lifcoiiH's  (K'tivc.  it  is  roticcivniiln  llint  if> 
■irti'iioics  tiii'.'lit  ililiiti'  lit  the  >;imii>  iiiotiKiit  tliat  tlioM-  of  a  iipiu'hl)orinL'. 

less   (it'tixc   cflltiT   li Mil-   <'iiM>t!iflril.       Alti'l'iit  imw    olivioiisl 


y  nii({ht 


••III'  nitlioiit  r!ni>iii!.'  liny  pi  riT|itililr  ctuiiiiic  ritliir  in  tin-  volume  of  tin 
Iniiin  as  a   w  lioli'  or  in  tlic  riiiiijition  of  xnioiis  (low. 

In  i-oiisiilciatioii  of  tlii'sf  t'ai'torH.  most  oliscrxcrs  art-  aurt'i'il  tliaf   tin 

'"'■'i'  \o|iiii (  III, Mill  in  tlir  liraiii  iiiiist  lie  roiistanf  at  all  tiiiH's  (Moiim 

ami  Ki'llii'  iloi'lrinr  .  Altiralioii  of  lilooij  supply  i-aii,  liowovcr,  still  \><- 
lin.UL'lit  aliiiut  li.\   rhaiii-'is  in  ||ii.  \rl.irity  with  wliirh  tin-  Mood  travcrsfv 

ilir  msmIs      WIiiii   •!■  liliMMJ  is  rfi|uiiri|   in  till'  lirain   to  supply  tlif 

inrri'asnl   Hii'talinlisni    wliirli    \\r   iiiiist    pii'sMiiii'  ; oiiipaiiii's   hfiuhfcncil 

iin-ntal  arli\il>.  it  is  not  ari'ouiplisht'il  a.-,  in  other  parts  of  the  hodv  1).\ 
an  inereasf  in  tin'  eaparity  of  the  vessrls  as  eonipared  with  tiioso  of 
other  \aseiilar  areas.  Imt  l.v  ;i  inirrviiiy:  up  ..f  (h|.  eireulatioii  throuu'li 
Vessels    \(  hose    ealilier    remains   uiialtereil. 

Till  iiniiii  fii.hiis  il  U  nil  ill  in  1 1  till  rihifilii  „f  hliii,<llli,w  tlirouijh  Hi, 
liniiii  iiiiisl,  till  n  Inn ,  hi  ih /ii  ml,  nl  iijutn  rliiini/oi  iiiiiirriiiii  rhcivhrn 
III  tl,,  r,is,'iiliir  sii^liiii.  a  eoiieliisii.ii  fur  whieh  there  is  ahundaiit  experi 
mental  e\ii|eiiee  (If  the  many  iii«eniiiiis  methmls  lliiit  have  hoen  de 
vised  to  seeiire  this  evidinei.  We  will  eite  hut  one  in  this  plaee.  Keponis 
are  taken  of  ehaiii^es  in:   i  I  i   the  r,  11,111^  tilnml  iinssiin    of  the  l)rain  h\ 

'■' li"K  •■!  <-iMinula  either  with  the  vein  immediatei.v    aft.'r  leaving' thi' 

•  kiiil  or.  Iietter  still,  with  the  Inniilnr  lli  miiliili ;  (2)   the  hniin  roliiiin. 

\,y  eoiineetinsr  a   very  sensitive   r 'ivin^'  Innihoiir  with  a   trephine  hole 

in  the  eranium  so  that  its  open  end  lies  auaiiisf  the  pia  mater.*  .\l 
tliiiU'-di.  as  ue  have  seen,  while  ineaseij  in  the  rijrid  eranium  fho  brain 
'Illume    can    not    ehanne    i.i    any    deirree,    \et    this    will    oeeiir    when    a 

pirtion    of    the   eranium    is    removed,    so    that     pulsations     •respond 

illL'    to    those     in     the    111 I     vessels     will     he    iihservi'd  ;     I :! )     the    eireula 

tory  eotiditions  elsewhere  in  the  hod,,  hy  takiiu,'  arteiial  ami 
venous  pressures  .•m,]  pleth.v  smoniaiiis.  The  results  in  a  normal  an 
iiiial    show     the    Inilowiiiir    points    i  see    I'ii;.    Si))-     (li    '|'||,.    traeiliKs    ol 

the   arterial    hi I    piessii:e    (.li.   the   hrain    volume    iC)    and   the   intra 

iranial  venous  pressure  U'l  have  e.\aelly  the  same  eontour  -that,  is. 
the  respiiatiiry  and  the  eardiae  waves  in  all  thice  of  them  are  ideiifieal 
The    venous   Mood    as    it    Ihiws    into   the   .juirular    veins   also    pulsates   in 

1  I  " '■■  ;'  ''.!''i''';'  '•  'r'l-'i  f-'ih  ■■■"-;-l-  "l'  -i  I'l.i-  eiiu-  ..f  ih.-  v.iirr  cli.nni't.T  ;is  the  trc-nhn  ■ 
Mr  into  «h..h  ,t  ,«  I.Khtly  Mlr.i  11,..  l„,i,.  ,„i„.  „  ,|„.,,|  ,„  ;„  j,„„,,  ^.,„j  (,,  ,|,|„  r„,,,^„  mcmWanr 
.<nil  il!,  ■.,ilcr  iri.l  n  imiiKil,.!  mili  tin-  rciivim;  laml.oiir. 


Kci  i.i,\niTii:s  "►'  iii.iMin  si  iti.v  in  tiktaiv  visikkv 


251 


iiiii<><iii  witli  iIk'  iirtorv.  I'Ji  Aii.\  ••liiiiiL'f  in  IIh'  IiIihhI  picHHurf  ni'  thi' 
-vslcinic  xi'iiiMis  s.xsli'iii  is  iiiiiiii'(|iiiti-ly  iTllccli'd  in  tlif  lilmiil  prcssuri' 
ul'  tlx'  siiitisfs  III'  the  lii'iiin  iiml  in  tin-  luain  Milunic  'nut   ucll  slmwn   in 

.11 nipiinyintt  trnrintji.     i^li    A   duinm'  nrviT  iiis  in   the  vcssfl.s  i>t" 

till'  liraiii   \Nliii-lt  <'ini   not    lir  iiri'mintt'il   for   liy  Mimr  I'luintrt'  m'<Mii'i'i!ii! 


lit  '•11  Tn  .I1..W  >iiniiltnnrims  rri'.i.N  -)  tin-  .:rtrri;il  Mr..-. I  prr^suri-  (.4>.  tin-  vtii..ii-  jii.  ^ 
•  i/'i,  'lie  inlra.  i.tnial  i.rr^^iirc  (i"',  tin-  i  r<  "s-urc  ii'  Ilti  \(i-..'i-  '•inn**-  i'''.  Tin-  till  iii  .it 
,1  |.ii.,iii<-  |.r...|iin'.l  liv  ,timul.iti..ii  ..I  III.  I'li.l.i.il  .Mil  .It  ill,  \.ii:  1-  v  ill  1'.  f'"iii.l  l-  •  ..  i-i 
I  itl  iif  mtra.  raiiial  .nn'  i.rcliTjil  v.ti-iii*  |>r.>^iii.-.  ;n .  nin]-aii>  iiiw  th.tl  in  Ilif  ardlit^.  t''it  a  li^" 
ili.it   .if  llif   Miiiiii-    sy-liiii.      I  rr.iiii    LiMiiaril    llill.i 

^-'ulicrc  in  tlic  Viisculiif  svstcni  miKiiU'  Ilif  rijinial  fii\ity.  This  vv 
lit  is  iriiport;int  liccanM'  it  sliiws  tinit  llicic  fan  nut  lie  \  a.M'ninl'ir 
'i'\c  ciititiiil   lit'  till'   lirain    \i'>si'ls. 

Takint,'  into  (■(iiisii|('i;itii'n  nut   only  tlic   ri'snits  ui'  sncJi  cxpfriiui'iiis, 
>it  jilsci  the  peculiar  piiysleal  (■nnditiiius  f.\i>sling  ii.  the  cfauiiil  eavity. 


I'i 


'    i.'il 


!  'I 


>   M 


11 


M^ 


Tin;  ciKci  i.ATiox  OF  Tin:  lii.oon 


\\r   iriiist    i-M,clii,|..   Iliiit   .•liMlii.'i'N   ill    1)1,„)(1   su|)l>ly   (Icpciid   on   cluniircs 


till'    \cli)cit\     i<\'   tile    1 


iliiiidildu,  iiiiil  tli.it   siicli  .-literal 


(Ifpcinlciit    upon    cliiiimi's  oi-ciiniiii.'   in    tlic   nortic   iiml 
'n  tlic  vciia-cna  incssmv.     Wlicn  tlii'  aortic  i)n'ssiin'  i 


il  lolls  ill  volocitv 


\v  ill  llnH    iiiid  the  ccifliral  art 


I'i'ics  aiKl  move  aloii 


more  especial l,\ 
ises,  more  lilooii 
tliciu  at  an  increasril 
velocity,  the  in.  rcascl  pressure  prohjiMy  eailsiiifr  j,  iiie,lerato  degrc, 
of  passive  (lilalatioii,  to  all 
■ill 


>\y    extra    room    for    wliieli    tlie    tiumer 


meroiis 


snioii  ccre!,ra|   veins  liceome  eompresscl.     Tliis  eonii.irssioii  of  tlie  veins 
I'lohaMy  ,|o,.s  not  ohstriict  tlie  -n...,ter  Mow  of  Mood  tiiroujfh  them,  h- 
cause,  taken  as  a  whole,  tlie.x   are  ordinarily  milch  more  capacious  tl 
need  he.     <iii  the  other  hand,  if  the  aortic  ))ressiire  should  remain  co 
slant,  'ml   that  in  the  \<iia  cava  iiicreas,-,  Ihen  there  would  he  ohstriic 

tion    to   the   i)as.sa-/e   >,:   M 1    in    the    intracranial    arteries,   and    conse 

'lUi'iilly  a  diminished  velocitv  of  How. 


laii 
II 


Vasomotor  Nerves 


It   III 


iMi-ht    he    inferivd    that,   sim-c   the    l.|or,dl|ow    tlironyh    I  he   cerel.ral 

vessels   is   mainly   de|)en.lent    on    vascular   i litioiis   elsewhere    in    Hm 

I'ody.  tli.re  would  he  no  need,  as  in  th,.  ves.sHs  of  other  va.scular  areas, 
l-^r  vasomotor  liheis.  Ilistoloyists  have,  however,  discovered  the  i)res 
<■!•'•  ol'  surh  lil.ers.  and   it   has  hecome  necessary   f.u-  tne  physiolo-ist  to 

"'"'  '""  '•'  ""■>   ■"•'■  i<'-'il'.^   "I'  iiiiportance  in  ( ection  with  the  rejiula 

tion  or  the  Mood  supply  I,,  the  hr.;in.  Kven  if  it  is  admitte.l  that"  the 
arleno|,.s  coiihl  not  contract  or  ex|)an.i  as  a  whole  without  produciiiL; 
l"cal  eh.in.-cs  in  venous  inessure  or  cranial  v.ilume,  it  is  yet  of  eours( 
always  possilde,  as  has  already  h.vn  i-ointed  out.  that  one  set  <d'  arte 
Holes  iiii.jjit  contract  at  the  same  moment  thai  another  set  expamlcd 
That  the  vessels  call  iind,'!'-..  a  process  of  eonstricliou  has  heen  sho«  n 
I'y  expciinunts  in  which  the  volume  of  oiitllow  from  the  vessels  oi 
the  l.raiii  was  measured  in  perlused  pi  eparaticms  of  hraiii.  WIhi, 
•piMcphrine    was   a.lded    to    the    perfusion    tlui.l,    eurtailm..i,t    „(   outllou 

"•"  "'"^'-'-v'''!  t. ur  (Wiii-crsi.     Since  this  drusr  causes  con.slriction  ol 

y'sscis  only  uheii  these  ;iiv  supplied  with  constrictor  fillers  (sec  l)a;;v 
"•"''•  ""■  <'""clusi,,i,  may  l,e  draun  that  the  cerel.ral  hlood  vessels  ,h 
.•ontaiii  such  n,.rvc  libers.  Xevcrtheless.  the  local  va.somolor  control  ol 
Ihc  cerel.ral  l.lo,,d  vessels  can  not  have  Hi,,  sinnidcance  in  connection 
with  chauu'es  in  Moo.l  supply  that   il   has  for  oiIi.m'  vascul.-ir  areas  HliU 

•'""'    ■^'•"•' '  ■  •'^'"    'I'"!''!    ii''i-v<'    lihers    are    j.rcsent    in    the    oerehra: 

l.h.od  vessels,  ;,iid  pr..siim;,|,|y  under  certain  conditions  they  nro  eapal.l. 
"t  cnusin-  tiir  Mn.Mi  vessris  to  ithd.-i -o  alleialioiis  in  caliher,  hut  it  i- 
iiupo.ssihl,.  /„  s.,   nf  irh.il  n.,1  vnliN    llns  run  h,    inuhr  uornuil  i,>u,liti„„s 


m 


■i:cii.i.\RiTii;s  OK  ni.ooi)  sipi'I.y  in  crHTAix  visciiu.v 


253 


Intracranial  Pressure 

One  word  inoro  with  rciranl  to  wliat  is  known  lis  iiitrdiriiiliil  pnst^iirf. 
lliiit  is,  tlic  incssun-  in  the  spai'i-  liftwccn  tlic  skull  and  tho  brain. 
I  iidcr  ordinary  conditions  it  must  lie  ciiual  to  that  in  tlip  perel)ral  ('ai)il- 
l.irics.  and  may  hv  mcasurod  Iiy  (•onn('<'tin^  a  sensitive  nianoiiieter  witli 
a  lulie  .screwed  into  the  cranium  as  (hscriiied  ahove.  It  lias  heen  found 
to  vary  from  0  mm,  ll-r  in  a  man  standins;  erect  to  .")(•-(;()  mm.  Hi:  in  a 
(loir  jxiisoned  liy  strychnine.  It  liecomes  in  reased,  not  only  liy  eom- 
IMission  of  the  veins  of  the  neck  and  l>y  an  increase  in  •lenoral  arterial 
pressure,  liiit  also  in  iiatholoij:ie  conditions,  such  as  hydroecphalns.  A 
new  firowlh  in  the  liraiii.  if  it  oi'cnpies  more  space  than  the  tissue  wliieh 
1^  ilestroycd,  CNcrts  ))rcssnre  on  all   parts  of  that   refiion  o|'  the  cranial 

■  ivity,  lint  this  pressure  may  not  he  traiismilled  eiiually  throiiirhont 
ilif  cranial  coiit<'nts.  for  the  falciform  liiraments  and  the  teiitoiiuni  siij)- 
I'Hit  a  i)art  of  it,  thus  dirccliii^  the  spread  of  piessure  alonir  certain 
p.ithways.  The  structures  at  the  li.-ise  of  the  hrain,  the  optii-  nerves, 
I  lie  veins  of  (!aien  and  the  Sylvian  aipieduel  are  most  afl'eeted  in  this 
\Miy.  If  the  ])rcssure  is  rapidly  ap|>Iied,  however,  it  may  rise  throu<,di- 
(iiit  the  cranial  contents.  In  such  cases  the  ]»ressuie  is,  of  course,  eir- 
I  iilatory  in  oriy-in,  since  immediately  after  death  from  c(>rehral  tumor 
the  intracranial  pressure  is  not  found  to  ho  I'aiseil. 

The  nia.jor  siinii)tonix  iif  riri  linil  ciiiniiri  ssioii  are  no  (hiulit  due  to 
.iniiiiia  of  the  medulla  oliloiifrata.  which  may  he  tho  result  cither  of 
pessiire  ajjplied  locally  in  the  liiilliar  reirion,  where  the  |>reseiiee  of  a 
'  •  ry  small   foreijrii   hoily  or  only  trivial   tumor  formation   is  sunic-ieiit   to 

■  it'-troy    life,   or   of   jiressuro    transmitted    from    the    eerelnal    ca\it.\',    in 

V  liicli  case,  on  a< unt  of  the  snpi>ort  olVercd  liy  the  leiitorium.  a  mueh 

l.iL'cr  trrowth  is  re<piired  to  alTcct  the  medulla.  Iiiti'rnal  hydro<'e|>Iialils 
i  Induced  liy  hlockiiiij:  of  the  aipuMliiet  of  Syl\  ins  and  the  veins  of  (ialeii 
',)iiscs  the  jrrejilest  rise  in  intracranial  tension,  and  may  alTect  the  iiie- 
'i  ilia,  lu'eause  the  liraiii  is  driven  downwards  s<i  as  to  ])in<'h  the  luilli 
.  jniiist  the  occipital  lioiie.  It  must  lie  emphasized  that  it  is  not  the 
I'^i'ssiirc  ixr  sr  that   causes   the   symptoms,   hut    the   attendant    anemia. 

■  I'  symptoms  of  acute  cereliral  anemia  and  of  compression  lieintr  i<!en- 
'mI  (l.eonai'd  Hill''').  To  i-elieve  the  com|)ressioii,  ticphinin^r  is  the 
•iHinoii   jiractice.     The  tre])hine  hole  should   he  as  lary:e   anil   as  near 

the  source  of  comjiression  (tumor,  efe."l  as  jiossihle. 


CIRCULATION  THROUGH  THE  LUNGS 

Hie  pulmonary  nc  lesser  circulation,  as  it   is  called,  is  <piite  dilTcrent 
'111  tho  systemic  cireldatiou.      In   the  first   iilace,  liecanse   the   |)ressiiie 


'Is   -'*-  P; 


f   it 


; 

' 

a 

,,1;: 

.t 

L':i4 


•nii;  ( iisci  i.ATrox  of  tiii:  iir.iMin 


ill  llie  imliiiiiiiary  ai-lciics  docs  not  aniouiit  to  nKirc  tliiiii  iil)oiit  ^(t 


Hir,  or  aliout   oiie-sixtli  ol'  lliat   nl'  tlic  sy.sicniic  artcrios,  the 


peri  1)1 


icrai 


resistance  in  tlie  lilnod  vessels  of  tlie  luiijis  is  iiiueli   less  than  that 
the  Iiody  in  <:eneral.     This  lower  resistanee  is  owiii};  jiartly  to  the  laii 
diameter  of  the  arteiioles  and   the  small   amount  of  mnseulnr  fd)eis 
their   walls,   and   jiartly   to   the    fad    that    the   eaiiillaiies   are   held    eo 


staiitly   in   a   somewhat    dihited 


eomlitioii    on    aceonnt    of   the   suhatmoN 


I'liiTie  pressiire  in  the  thorax   i  s.m'  |iaf.'i'  .'iddi. 

Another  i)eenliaiily  of  the  ])nInionary  eirenlatioii  is  that  the  ealihei 
of  tlie    vessels   is   to   a    very    larj>e   extent    dependent    npon    the   ehalij;!- 

''i"t   '"'  ill   the   intralhcn'aeie  jnessiire   with   eaeh    inspiration   and   e\ 

liiration.       They    heeonu'    dilated    on    inspiration    and    eontraeted    on    e\ 
l)iratioii.      The    extent    to    which    these    respiratory    clianu'es    alTect    iIm 
amount   of  Mood  ccjutaincd   in   the  Innirs,   is  very  considerable.     At   tin- 
heiirht   of  iiisjiiratioii    it    is  cnminited   that   a    litth'   more  than  eii-ht   Jici 
cent  (d'  the  wloile  hlooil  in  the  liody  is  emitained  in  the  hin!,'s.  whereav 

\I)iration   it   dimiiiisjics  to  lictwecn   live  and  seven  ])cr  cent. 

A  third  peciiliaiity  is  that  the  pnlinonic  Mood  vessels  are  not  snp 
lilied  with  vasomotor  nerve  lil.ers  at  least  with  snch  as  can  readily  1.. 
demonstrated,  II  is  said  that,  when  the  pnlnionary  vessels  are  per 
fused  and  the  cnittlow  measured,  a  diiiiinntion  in  the  latter  is  found  to 
occur  when  epinephrine  is  added  to  the  in.jeetion  flnid--a  result  whieli 
is,  liowever.  denied  Iiy  certain   investiirators,     Chanjies  in  the  liloodtlov'. 

have  not  lieen  ot.served  t ■eiir  when  the  vau'iis  or  sympathelie  nerv. 

tihers  runninir  to  the  Innjrs  ;ire  stimulated.  In  short,  the  coiudusioii 
whicli  we  must  draw  is  niindi  the  same  as  that  for  the  lilood  vesscK 
of  the  hrain  namely,  that  althouuh,  as  ,i  result  of  the  epinephrine  e\ 
l)erinienf,  we  must  admit  that  ;i  viismnotoi'  siipi)!y  iinni  ])o.ssililv  Im 
Iiresent.  yet  it  is  on<  whi.-li  can  U,'  of  no  sifjuilicance  under  inn'ma' 
conditions. 

When  there  is  olistrudion  to  the  ontllow  of  l.lood  from  the  left  veii 
tricle.  as,  foi'  example,  in  cases  of  hiyh  aortic  |nessnie.  the  Mood  is  n..- 
entirely  <lis(diaiire.l   with   ea.-h  heat   of  the  l,.f|    ventricle,  ;iiid   therefor. 

dams  hack  thiouuh  th.^  left   anriele  into  the  lun-fs.     On  ! oiint  of  tli. 

marked  distensil.ilily  of  the  imlmonary  capillaries,  ii  larire  amount  o 
this  Mood  may  collect  there  and  thus  m.ikc  the  Innu's  serve  as  a  kind  '• 
reservoir  of  tin-  heait.  When  the  capacity  of  this  reservoir  has,  how 
ever,  been  oversti>i>pcd.  an  increjised  iicripheral  resistanee  will  come  I' 
lie  olTercc]  to  the  movement  of  Mood  iu  the  pulmonary  arteries,  th 
pressure  in  which  will  consc(|Uently  rise  and  sooner  ui  later  interfer 
witli  the  discli;ui:e  ffom  the  riirlif  ventricle,  causing  as  a  result  a  staii 
nati(ni  of  blood  in  the  systemic  veins,  and  a  eonsc(pn'nt  increase  in  vo! 


ri-,(M  i,iM(iTii;s  op  iti.oiii)  sci'i'r.Y  i\  cutTAiv  visci:i!\  2;);> 

uiiic  of  such  viscoru  iis  tlio  liver  aiitl  kidneys.  Tlio  siinie  changes  will 
nliviniisiy  also  supervciio  wlicn  there  is  ref^urj^itiitioii  of  lilood  fi'oiii  tiie 
It  It  veiiti-icio  1o  the  left  auricle,  as  in  eases  of  iiiiti'al  iiisuflicieiiey. 


CIRCULATION  THROUGH  THE  LIVER 

The  liver  is  the  only  irland  in  the  hoily  reeeivinf;  hoth  \enous  and 
arterial  lilood,  the  hirnier  lieinj;  supplied  to  it  at  a  very  low  ])i'essure 
liy  way  of  the  eajjacious  iiortal  vein,  and  the  latter  at  very  liiirh  pressure 
liy  the  strikinK:ly  narrow  lu'i)atie  art(>ry.  Except  fiU'  the  i'elativel.\- 
Miiall  amount  of  l)lood  which  is  su]>|)lied  to  the  walls  of  the  hlood  vessels 
.lud  the  liiliary  ducts,  none  of  the  lie])atic  aitery  lilood  mixes  witli  that  of 
till'  iHirtiil  vein  until  the  vessels  enter  the  hepatic  lohules.  I'eyoinl  this 
|i.iiiit  the  two  lilood  streams  mix  and  tlii'  comliined  sticam  is  drained 
:\\\:\y  ]ty  the  suhlolmlar  and  hejiatie  \t'ins. 


ill 


Methods  of  Investigation 

To  study  the  relative  imjiortanee  of  these  two  sources  of  lilood  sup- 
lily,  and  also  to  iiivestif;ate  the  manner  in  which  the  latter  is  controlled. 
the  iiH)st  sntisfactoiy  methoi!  has  consisted  in  mcasurenwiits  of  eluinjres 
ill  volunio  flow  ratJu'f  than  in  those  of  clian>res  in  jiressure.  The  v<il- 
UMie-flow  measurement  has  lieen  made  eitiiei'  liy  eoiuiectiuir  stromidiis 
liaire  20")  to  the  hej)atie  aitery  or  jxirtal  vein,  or  hy  measuiiii};  the  <iut- 
I'nw  of  hlood  from  the  hepatic  vein  into  tlie  \cna  cava,  first  with  liotli 
iiillow  vessels  intact,  and  then  with  one  of  them  lijiated.  An  oli.jec- 
'inn  to  the  first  (the  stromuhr'i  method  is  tin'  jxissilile  interference  with 
i'liiodtlow  or  IiIikmI  pressure  prinluccd  li.\  insert  iiijr  the  stionuihr  into 
till'  eiiteriufr  \csscls.  and  also  the  fact  that  simultaneous  nu-asuremeut 
'■!'  the  flow  in  lioth  vessels  can  not   lie  made  satisfaetoril.v. 

To  measure  the  outtlow   from  the  liepatit-   veins,  the  aorta   is  lijjated 
'■'low  tlie  celiac  axis  and  a   ^^  idi'   i-aiinula   is   inserted   into  the  central 
'lid  of  the  vena  cava  liehnv   the  levi'l  of  the  liver,  a  loose  thread  licinir 
"i.iced  ai-ound  this  vessel  just  aliove  the  ilia|ihratrm.     I'.y  pulliii<r  on  this 
'iiread    the    vena    cava    licconn-s    oliliterafed,    and    the    lilood    from    the 
' 'patic  veins  is  therefore  diverted  into  the  cannula,  throiiu'li  which  it 
iws  into  one  end  of  a  \t'ssel  shaped  soniewhat  like  a  sputum  cup  (tin' 
!   'I'iveri,  the  other  end   lieinv'  connected   liy   tuhiiii.'  with   a   piston    re- 
order, from  tin-  movement   of  whicli  the  volunn-  of  lilood  flowiiif;  into 
lie  receiver  can   readily  lie  computed.     To  nuvisure  the  thiw  of  hlood. 
■<<  clip  on  tlu'  tulie  of  the  receiver  is  removed  at  the  same  moment  that 
'iie  tlu'ead  ai'oiind   the   vena    cava   aliove   the   diaiihrasrm    is   tightened, 
■id  when  the  receiver  has  filled  with  lilood,  tliis  thread  is  ;i":aiii  loosened 


iri 


i    t 


■is     -      i 


.>."•(; 


THE   (  IKCI  I.ATIdN    OK   TlIK    JlI.CMU) 


and  till-  ri'ccivcr  lilted  ii])  so  tlial  tlic  lilixid  Hows  at  low  ju'cssurc  Iiin' 
into  tlic  fiiriilatioii.     'rill'  iTccivcr  licinir  »(  known  caiiacity,  the  Ifiinlii 

of  tiiiif  it  takes  till'  111 1  to  till  it  as  dt'ti'ii'iincd  liy  tlic  piston  record,;, 

riiiiiislies  ns  with  the  neeessaiy  data   fii'iii  which  to  calculate  the  ''at 
of  How.     The  feccivcr  is  chosen  of  siicii  a  si/c  that  it  takes  only  a  fe\ 
seconds  to  lill,  the  di\eision  of  lilood  into  it  not  causinj;  any  niateri;ii 
fall   in  aiteiial   pressure.     The  ohservations  ai'e  repeated  fre(Hiently. 

Results,  r.y  the  use  of  the  c  methods  it  has  Iieen  found  that  the  tol.i 
mass  iiioM'HU'iit  of  Mood  to  the  livci'  of  the  dojir  vai'ies  hetween  1.4(i  aihl 
Il.4(l  c.c.  jier  second  for  1(10  ^'lams  of  liver.  Conslderahle  ehaiijres  uv\} 
occur  in  the  arterial  iircssiire  without  affecliri};  the  liver  tlow.  WIhm 
the  hepatic  artery  is  occluded,  thi'  How  diminishes  hy  aliout  'M)  ]ii' 
cent,  or  conversely,  when  the  jtortal  vein  is  ohst meted  hut  the  hcp.-iti- 
artcr.v  left  intact,  hy  ahout  liO  per  cent.  iiidicutin<;  that  ahoiit  one-thii'i 
of  the  tot;il  liloodllow  thiouyh  the  liver  is  eontrihuted  hy  the  hepati'- 
artcr,\-  and  two-thiids  hy  the  portal  vein.  Sonn-  hlood.  however,  u:aiii~ 
the  liver  Ihroiiirh  aii.'istomotie  channels  hetwceii  it  and  the  diaphra'.' 
malii-  \t'iiis. 

TIk  nhitiri  siii>i)l!i  hi/  Ihr  lira  rtxnth  is  suh.jeet  to  various  eondi 
lions.  That  thnniirh  the  hepatic  .-irtery,  for  example,  may  he  very  con 
sidciahly  altered  on  account  of  vasoeousti'iction  in  this  vessel,  for  it-- 
\\:\\\s  can  easily  he  shown  to  he  liherally  su]>plied  with  vasoconstrict"' 
(ihers  cariied  hy  the  hepatic  ])lexus.  This  can  he  denunistrated  h 
the  rise  in  Mood  jiressurc  which  occui's  in  a  hranch  of  the  heiiatie  artei  \ 
duriiii:  stimiil.-ition  of  the  jilexus.     On  the  other  liand,  alterations  in  tli- 

hi IMow    in   the   portal   \cin   can    not    he   hrou<;ht   alio\it   hy   active   cm 

striction  or  dilatation  of  the  intralic])atic  hianchcs  of  this  vessel,  n 
aetive  \asomotor  (ihers  haxiiiy:  heen  demonstrated  hy  stimulation  .. 
the  hep.itic  ner\('s,  although,  as  in  the  case  of  the  lirain  and  luii^  IiIoim 
vcNsc'-  I  rt;iiii  amount  of  consti'iction  may  occur  under  the  intlncMi 
of  cpi,         .  inc. 

The  hloodllow  throuudi  the  jioital  vein  is  deix'iideiit  on  chaiifrf^s  m 
currin!,'  at  either  end  of  the  distrihution  of  the  vessel,  that  is,  chaniji 
occuiiint'  in  the  liver  itself  or  i?i  the  inti'stine.  Of  these  factors  the  l,i' 
ter  is  Mo  cloitht  the  more  imiMirt.int.  an  increase  not  onl.v  in  poi'tal  hhio 
jn'cssui'c  hut  also  in  jmrtal  hloodllow  heiiiir  reailil.v  jirodueed  hy  dil; 
talion  of  the  s|ilanclinic  hlood  vessels;  for  examiile,  as  the  I'esult  of  sc 
tion  of  the  s|)lanchnic  nerve.  Alterations  in  ])ortal  hloodHow  l)roii>.'l 
.-iliout  hy  chantres  in  the  ealiher  of  the  vessels  in  the  liver  itself  a' 
p.'iilly  di'pcndi'iit  ii)>o!!  cli.iniri'^;  in  the  hraiiches  of  the  hopatie  arte?- 
Let  lis  considei'  hricfly  how  this  ina.v  he  hi'otisrht  ahout.  At  the  poii  ■ 
where  the  ])oital  and  hepatic  arteries  come  together— that  is,  at  the  ii 


3 


i'i;<  TT.iAKirirs  of  I'.i.ook  m  i'm.v   in;  ciktain   sis(i;i!a  '2-u 

tiiilicpafic  I'iipilliiiii's  tlic  i)ii'ssiii('  lit'  tlic  Miiiicl  ill  flicin  must  Itccoiii"' 
(i|iuil,  \\iiifli  mi-iiiis  tli;it  In  its  cduisc  llircniuli  tin'  intcrlnliiilju'  coiuici'- 
ii\c  tissue,  flic  lii'.-iiiclii's  cit'  tlic  iic'ii.ilii-  .•iiti'i-y  iinist  otlVr  imu'li  resist. -mi't' 
to  llic  I)1(i<m1  llciwiiit;  tliiiiuiili  tliciii.  This  t'iictii)u;il  ri'sistiincc  icsiilcs  in 
tlic  Iicpatic  aitcrioii's,  iiiid  siiicc  tlii'se  jiic  riclily  sujiplii'.!  witli  cunstiic- 
iiir  iici'vcs,  ffi'cat  varialinu  in  in'iiatii-  iiillow  liccntni's  imssililc.  Tlicsc 
lii.intros  will  aftVct  tlic  (lc<!:rcc  nt'  tciisiun  ut"  tlic  intciloliuiar  coiiucctivc 
tissue  ill  ■wliich  the  arterinles  lie.  In  this  tissui',  linwcvcr.  also  lie  1he 
lliin-walied  bl'aiiehes  uf  the  iinrtal  vein.  When  Ihererore  the  feiisinii 
lif  this  tissue  lieediiics  irreater,  as  a  result,  tor  example,  of  vasodilatation 

in   the  liei)atie  artciy,   the   portal   vein   ladieles  will    Ik nu mpiessed 

iiikI  the  liloodliow  aloiitr  them  impeded.  ('mivciscly,  when  vasocoii- 
sliii-tioii  oceriis  in  the  hepatic  arteries,  the  eoiim'stion  of  the  coinifclivc 
ll^^llc  licconics  (1  liii  1 11  islicil,  tlic  vcins  dilate,  and  the  liloocl  flows  thioiiudi 
Ihi'iii  more  readil,\  ( .Xfaclcod  and  K*.  <i'.  I'carcc-' i .  Kxpcriiiicnial  e\  i 
'I.  ncc  in  sii])]iort  of  the  aliove  view  is  I'uinislied  hy  olisi'r\  iiiy;  the  oiit- 
llnw  of  hlood  I'roiii  the  liver  lieforc  and  diiriin;'  sliiniilalioii  o|'  the  hi' 
i'itic  ple.\us.  The  (irst  clTcct  is  an  increase  in  the  outllow.  which  \(r\ 
-•tpnii  retui'iis  to  its  oriirinal  amount,  even  thoii'.di  the  stiniulation  of  the 
pl'Xiis  is  kei)t  up  dliriiiir  the  c\|)eiiiiH'nt.  This  return  to  the  normal 
i' iw  must  indicate  either  that  the  constriction  of  the  hepatic  artery  has 
I  "t  lii'cii  iiiaintained,  or  that  it  has  hccn  maintained  Init  is  accompanied 
!■>  a  compeiisatory  increase  in  the  (low  throii<_'h  the  |>ortal  vein.  As 
matter  of  fact,  we  know  that    llie  Iicpatii-  ;iit','ry    remains  constricted 

■  -  iipiiir  as  the  hepatic  i)le\iis  is  stimulated,  indicitinsr  that    the  contres 

'  'II  of  the  eoimeetive  tissue  in  which  the  \enules  lie  has  heeoine  reduced 
'  --ueh  an  extent,  as  a  result  of  the  const  rict 'on,  that  these  open  up  and 
;  '  iiiiit  the  lilood  to  llow  Ihroiieh  them  more  readily.  The  initial  in- 
'  lase  in  outllow  immediately  follow  inn-  upon  si  iinnlation  ,,\'  the  hepatic 
|'''\us,  is  no  dolllit  caused  liy  the  s(|iieeziii'j-  out  of  the  Mood  alleadv  ill 
';.e  hepjitic  vessels,  and   it    is  a    result    which    is  often   oliservcd    in    otl 

■  '^ans  diiriuir  stiniulation   of   vasoconstrictor   nerve   lihers. 

THE  CORONARY  CIRCULATION 

We  have  .-ilready  studied  the  elTeet   produced  on   the  heaitlieat    liv    in- 

■  I  ieriii<r  v\  ith  the  flow  of  lil I  in  th irnnary  vessels,  ;ind  it   remains 

■r  us  to  study:  (ll  peculiarities  in  the  hloodllovv  thioii'^di  theiii.  and 
-  whether  this  liloodliow  can  he  altered  lp,v  dilatation  or  constriction 
■   'he  vessels  hrolifihl   ;ilititll   Ihroit-h  lur.  rs.     With   leuaid  to  liie  /<'..( 

iriliix  (if  blomlltoir,  it  may  lie  stal.'d  thai  there  is  said  to  he  two  periods 
11  each  cardiac  cycle  during'  v\hicli  mi   increase  lakes  pl.iee  in   the  m.iss 


I  1 


I 


fl';jii^    ^ 


-Li 


i>r)8 


■niK  ( iKci  r.ATiox  OK  Tin;  iii.oon 


inovenipiit  of  bluod  in  the  fiiioiiary  v.■^st'ls  iiaincly,  at  tlie  l)i'j;iiiiiiiiv 
of  systole,  and  ajtaiii  at  tin'  l)Oi;iiiiiinj;  ol'  diastole.  Nevertheless  tli. 
pressure  pulse  lias  the  same  eontuiir  in  the  eonninry  as  in  the  systemic 
eireiilation.  ( W.  T.  Porter.-i  Durini:  systole  the  intramural  liraneh.'^ 
of  the  eoroiiary  artery  are  compressed  and  the  Idood  pressed  out  i<\ 
them.  This  enii)tyin!;  of  the  vessels  favors  the  How  of  hloiid  throu>;li 
the  heart  walls. 

Reirardinj.'  the  i)resence  of  roroimrn  viisumolnr  lurvt.i,  tliere  is  at  i)res 
ent  a  eertain  amount  of  douht.  When  strips  of  the  eoroiuiry  artery  niv 
suspended  in  a  solution  of  epine])hrine,  they  uuderu'o  relaxation  instea'l 
of  eontraetioii.  On  the  assumption  that  the  aetion  of  ei>inephrine  on 
l)lood  vessels  is  the  same  as  that  of  cumulation  of  the  vasoeonstrietoi 
tihers,  this  lesidt  has  heen  taUen  as  evidence  of  the  altseiu-e  of  sneli 
fillers  and  the  jxissihle  presence  of  vasodilator  fihers.  A  somewhat 
similar  tvjic  of  exiieriment  has  lieen  ])erfonned  hy  injectinc;  epiuepli 
rine  into  the  fluid  tiscd  to  jierfuse  the  excised  mammalian  heart, 
with  the  result  that,  when  such  in.jeetions  aie  made  into  a  heart  that 

is  not  beatinir.  <'\  ideiu-e  of  \as nstriction   is  ohtained,  whereas  when 

in.jeete.1  into  a  heatini;  heart,  dilafati(.n  occurs.  This  latter  re.sull 
may,  liowever,  be  owin<r  to  the  action  ul  the  ei>ineplirine  iu  stiniulatin'.: 
the  eardiae  eontrietioiis.  Other  oh.servers.  however,  deny  that  the  in 
.jeetion  of  eidneiihrine  into  the  eoroiiary  circulation  has  any  influence 
upon  the  outflow  of  the  perfusion  fluid.  Takint:  the  result  of  these 
observations  as  a  whole,  we  may  at  least  eouehule  that  eiiinephriiie 
does  not  jjroduee  the  same  marked  vasoeonstricf ion  that  it  lu'odiiees  in 
other  blood  vessels-  a  fact,  which,  as  already  .stated,  may  be  taken 
ailvantasic  of  in  brinu'in!.'  about  the  rise  in  coronary  pressure  that  i^ 
lU.es.sary  foi-  sticcessfnl  I'esnscitaf ion  of  the  lieart. 

.'ttenipts  to  demonstiMte  llie  iiresence  of  vasomotor  fibers  by  cleetriiMl 
.stimuf.uion  of  the  vauus  or  sympathetic  nerve  have  yielded  results  Mhii'l: 
are  fpiite  iiu'onehisive.  although  some  observers  assert  that  the  vairu- 
nerve  carries  vasoeoiistrictor  fibers  to  the  coronary  vessels,  and  thai 
the  sympathetie  e.-irries  vasodilator. 


niAl'TKU  XXIX 


CLIXICAI.  AI'I'I.ICATIONS  (»F  CKFfTAIN  niYSIOLOGIC 
.MKTIIODS* 

III  flic  lollduiii!,'  cliaiitiis  a  liiiff  a xiiit  will  l.c  (.iTcic.l  of  llie  cliiiifal 

(ISO  of  liio  (•icctrocariliourain.  of  polyspliv  jrninyiains,  aii<l  of  Moodflow 
iiifasiiiciiiciits.  Tills  is  (loiif  to  sliow  liow  pliysioloijic  tci-liiiic  is  iieiiiij; 
rinploycd   for  the  accurate  iiivcsti<;ation  of  cardiovascular  disease. 


i  Ij^ 


ELECTROCARDIOGRAMS 

To  ohsorve  the  electrical  ehaiif,'e  produecd  Iiy  the  spread  of  the  exeita- 
lioii  wave  over  the  heart  from  auricles  to  ventricles,  it  is  not  necessary 
In  place  the  electrodes  directly  on  the  heart,  hut,  as  already  hinted,  we 
may  follow  the  electrical  cliansie  hy  h-ading  oiT  from  electrodes  applied 
111  the  surface  of  the  liody.  From  such  elect roeardiofrrajihic  tracin}»s 
extremely  important  facts  concernins;  the  projiafration  of  the  heartheat 
may  be  ascertained.  In  order  to  make  an  ohservalioii  the  iiaiids  and  the 
icfl  foot  arc  each  iilaeed  in  a  solution  of  sodium  cliloiide  contained  in 
|Mirous  jars,  illllllcl•^ed  in  Inryii'  vcs,scls  coiitainin;.'  a  saturated  solution  of 
/iiSO,  and  zinc  tcrminals.f  .\ii  arrainicincnt  like  that  in  Fijr.  SI  may  al.so 
Ik  used.  I'.y  maniiniljiliiin  of  siiilaiile  keys  the  extremities  may  then  he 
iniinecled  with  the  electrocanliotrraiih  in  the  followiiii'-  manner:  Lead  I. 
liu-lit  arm  and  left  arm:  le.nl  2,  rijjht  arm  and  left  leir:  lead  :!,  left  arm 
:iiiil  left  h'H'.     Tlirouu'h  lead  I,  the  current  actin-;  on  the  <ral\anometer  will 

i"'  that  pi'odu I  more  cspcciailv   ,it  liic  liase  of  the  heait.     Tiiioujih  lead 

-.  the  current  will  pass  thidn<ih  the  lonu;  ,i\is  of  the  he.iit.  and  throuirh 
i'lid  I!,  it  will  piiss    .lainly  aloiif^  its  lell   iiordei-. 

When  any  pair  of  leads  is  connected  with  the  jralvaiiomcter,  it  is  oli- 
served  that  the  strin>;  is  dellected  to  one  side  owinjj;  to  electrical  cur- 
rents arisiiijr  from  tlie  skin.  I'.efore  takin^r  a  record  of  the  cardiac 
riiovemonts  of  the  striiiir.  it  is  necessary  to  rompfiisatti  for  this  skin  cur- 
rent  hy   iiitroduciiiir  into  the  circuit    in   the   op)M>site   direction   the   re- 

•.\  <''rl.iin  ,im. .11111  .if  n  |..lili..u  ..(  mjlli  r  iirt\  i..u-ly  lii-i  ii--r.|  li.i^  lii-ti  fuun.l  .-i.lvi.il, I,-  in  ilu-se 
lu.|)ter.s  fur  wliiih   Ihc  iti(liiis»'iu-r  of  tin-  itiulrr  i>   rcnu-.lf.l. 

til  is  roally  ininrcrs-.Try  1..  \im-  llu-  sf.  i-.ill.  .1  n..ni..il;iri?.il.l.-  iliitr...lv«  C.l.i.i  v.-<.l.  c-..iil.iiiiiiiu 
•1  f.er  cent    \aCI    snliiti.m   willi   111.'   7iiK'   |.l.il.  s   .lii.i.ini;   iiil..    Ili.in    an-   .|iiit,-   -alisf.iclory. 


il 


hi  > . 


i  ii 


2(10 


Tin;   (  llti  I  l.\Tln\    or    iiri;    I'.l.iiiil) 


!■     II 


y 


I:   4 


i|ilirfi|  ,'iiiiniiiil  III'  riini'iit,  rallfil  tlir  (•<.iii|>ciisal iiii;  riirrfiit.  to  lifiiii;  lli< 
Ntniiiy:  sli.iiinu    liiick  tn  tile  /do  or  III i<l|>nsjt jdii.     In  iiiil.'f  Hint   tlio  n  , 

tivf   ill   clijiiacliT,   it    is   I'liitlu'i'   ncccssiH' 
lie  sill  11(1, iril'r.iil.     'I'liis   is  ilciMc   liv   iis 


"111  iilptiiiliccl  may  lie  (|iiaiitilat 
tlial  till'  iiHP\  I'liiciit  iif  tile  stiiii 
ci'ilaiiiiiiu   l<i   uhat    cxliiil    llir  siiinjr   mnvrs   when   ii   ciiiri'iil    i.f   1, 


\\>i\\  I 


illaL.'!'  is  sent  ||iimiil;|i  it  ami  li\  aiti'riii<:  tlic  ti 


llsillll    III'   ihl'   stlil|i_r   Sll    til 


"illc    liliHiMill    111'    iMIIIrllt 


c-ausis    an    CNrlllsKHI    nl'    mii'    rrntiliiclcr    nl'    tl 


striiii:  sliaijiiw  mi  tin'  |iiii.lnt;ia|iliic  plat.'.      It    uuiilil  tak.'   as  lii-viiiiil  tl 


'irilil    jn   till-   iniitt  \1    ;iri    pri  li  KiM. 


ciiiidiii's  ..r  this  Miliiiiic  tu  Mil  ill  any  aiTaliT  ilctail  into  tlic  tccliiiic  in 
volvcil  in  takiii-  I'ln-t  rm-anliniriaiMs.  Init  it  may  In'  saiil  that  tliis  is  1.. 
no  miaiis  .lit'linilt,  piux  iilcl  tlic  iiisl  niftions  \\  lijcji  ai'c  MippliiMl  wili 
tlir  iiisliniiiint  an-  i-aivfnjjy  roll, ,«,.,].  In  i)racti,-c  tlio  1akin<r  of  I'liv 
trocai'ilinmaiiis  is  imli'i'il  i|iiiti.  a  siiM|ili'  malliT.  ami  tlii'  cxtrciin'ly  im 
imrtalit  inl'oiinaliiiii  whirli  tlu'v  uivc  us  coiic'i-niii^'  the  iin'clialiisiii  i. 
\\\v  ju.arlli.'al  ami  tlif  f.  iilrm-r  ..i' 


I'liiployiiiriit   a   iin 
•I 


IliyocHi-ilia!   ilirM-asc  simtiii!   iiiakr   th 


iivfisal    piai-tii'c   in   all    caiiliai-  el 


inii-s.     Siiiiii'  of  til 


cliiiK-al  applications  aiv  ili  si-rilinl  ciscu  liciv  (  pau'c  l.'(i(l 


I  I.I  r'j'liiir\lii>liN;i| AMS 


.'III 


Wliiil  piiilKiiliirly  iiilricsts  IIS  licit'  is  ill 


iliitir  III   llii    I  li  iliiiriinlii 


•  ii-iiiii   III  II   iiiiiiiiiil  jii  i-siiii    I  I-'mj.  S'_»'.      ll    will    111'  (iliM'iM'cl   thill    Ihi'ic  iin 


I    I 


l|     lUll     WIlVl'S     llt'id 


w   it. 


iliic'i'   w.'iMs  iilicivi-  till'   line  111'  z,.|ii   piili'iitiiil    nil 

TIk'.v    li.ixc    lictii    Ictli'iiil    riniii    lii'lurc    liMikujiiil.    I",   <^,    |{.    S.    iiinl    T, 


ll     ill    .'ill     slll'l 


I    iiTiinls    «  hi'li    riiitri't  l\    iihl.-iiii 


'I,    ll 


11'      V\ll\l's     mIiiiVI'     till 


IMC   111'  /ciii    |)iilciili;il    iiiilii  iitc   til,, I    ll 


ll 


IC    tljll 


'\.    T 


IC    C\iicl     C-nisc    111'    I'JH'll     W  il\  1 


li;isc    111    llic    licjiti    IS    iiclmIi\c   1i 
lills  I 


c    lllls    llci  n    ,'lsc(|t;|lllci|    1)\-    tllkiM 


-iiiiiiltiiiiciiiisly    uilli    till'   il(i'triii-;ii-i|iii'.'f;ii 


;iiici's  iicciiniii'j  III  tl 


rl   ll 


IC   licMI'l    illlllll'.'   I'.lcli   c;iri|i;ic   c\  cji 


11     il      lll'lllll     111      tlic     III 


CCllillllCllI 

Sllcii    iccuiils 


1 


f       T 


I 


I  i-.    .'•-■.      .\..|.i:.l    .l.'Ii..,:,r,!i..LT.iii:        I.,.,,;,    1,    J,    ,!,       \,,,,     i|,,„    ,1,,,    |,.  jj,,    ,,,    ,;„     /^    ,|,  i), ,  ,,,,,,    ,,, 
l'"l    ■*    <inil-    111,-     lill,i,iii,     l„i.,,,.„    i|.;.    ;„,i,.|,|    ,.r    /.,',    ,,,.,1    ,v^. 

!i:i\c  liccii  scciircil  liy  t.ikiiiy;  iiilriic'irdiiic  prcssui-c  cuivcs  with  till'  results 
:>  shiiwii  ill  Fiu'.  S'l.     Tlic  tup  ciiivc  icpicsi  iits  iiiiriciiliir  iind  the  sccuml 

•'lie    vciitriciiliii-    pressure,    whereiis    the    lowest     is    ;iii    elect  r((c;irirni<;r,'nii. 

It  will  he  iihservid:    (\\   tluit  the  I'-wiive  necurs  Just   jiiiteeeileiit  tii  eiui- 

•I'.M-tiiiii  of  the  luiriek's:  rl\  thiit  the  siiuill  pusitive  ujive.  (^,  u  liich  is  ah- 
'lit  ill  these  traeilifis,  must  (leeiir  Just  Ixliire  the  lii'!:iiiirui<.'  ol'  the  ciiii- 
I'iictiim  of  the  ventricles;  (I!)  that  tlu'  iicirative  wave,  J{.  ciceurs  just  lie- 
•re  and  duriui,'  the  early  p;irl  dt'  ventricular  .systi.le — that  is,  during; 
lie  prcsphyfriiiie  jieridd:  and  (4  I  that  the  loii!,'  ujiwiird  wave,  T,  euliiii- 
•  ites  at  the  iiKiiiieiit   the  xcntricle  lieijins  relaxiiiir. 


%i^M 


>   ,. 


1 1 


.'62 


Tin:    (  IKII  I.ATION    OF   Till;    lll,(Mii> 


Altliouuli  siirli  rniiip.iiisiiiis  ifivc  us  «M.ii>i.|,.,al)l("  iiiNiKlit  into  flic  cause 
of  several  nf  the  ^\a\es,  tlicre  vet  leiiKiiii  certain  peculiarities  of  tin' 
eleetiocaidiotrriiiii  to  lie  eonsidereij.  Tlioe  are:  ili  the  cause  of  the 
slit'llt  I)ositi\e  wave.  (}.  i  •_»  the  cause  of  the  positive  wave.  S;  CJ)  the 
i-aiise  for  the  pcrio.l  of  .•.niai  poinitjal  at  the  liase  and  apex  durinu'  \en 

triciiiar  systol.-  indicated  li.\    the  port! f  the  curve  lietwcen  S  and  T; 

'■*^   '• ''"1^"'  •"•"■  llie  ne!.'ative  wave.  T.     To  solve  these  iiroMcnis   it   is 

necessary  t niparc  electrocardionrains  taken   from  flic  surfa f  the 

l)ody  witli  thos,.   froni  electrodes  placed  diieell.v    on  the  l.ase  or  apex  of 
the  ventricle  of  tl xpoM'd  iii'art, 


ll.^ 


i 

1  If. 

L 

j..-t    I..  |..i,>    [],.■    |.i,-|.|nK„n,     i„,,.,l    M.iit-    111    Hi,     »,i,tri.  I.  .      (!■>.. ii;    I.,  im-.  I 

The  Ventricular  Complex 

In  view  of  the  nature  of  the  electric  chanu'e  which  occurs  in   a  strip 
of  (leiicrvated  muscle  when  a  wave  of  contraction  passes  alons;  it   (pai:<' 
ISSI,    till-   siiiiph'st    inteipietation    of    the   ventricular    part    of   the   ahov.- 
curve  is  that  the  contraction  must   pass  into  the  ventrielp  at  a  little  dis 
tance  from  the  liasc,  thus  eausini,'  the  latter,  for  a  moment  of  time,  to  he 
positive  to  the  rest  of  the  ventricle,  :ind  aceountinj,'  for  the  sli;jlif  down 
Avard  wave.  {}.     Immediately  after  this  the  hase  of  the  ventricle  liecomcs 
nesrative   to   the   apex,    «ivinir   ns  the    marked    upward    wave,    1{.   wiiich 
however  lasts  for  hut  a  short  period  of  time,  lieintr  followed  I)y  an  inter 
val  liurintr  which  the  hase  and  ap.'x  are  of  the  same  electrical   potential 
( horizontal  i)art  of  wave  between  |{  and  T).     I'inally  the  li.-ise  atraiii  be- 
comes nc<;ative  to   the  ajiex.    tiius   aecountiiij;    for   the  smaller    upwai'd 


KI.K('TK()I'AKI)|)NIK.\MH 


263 


uiivc,  T.  Tho  vtiUHv  (if  the  iicciisi<miill>'  (iliscrvfil  iIiiwiiwhi'iI  wave,  S, 
fcllnwinu  I{.  is  olisi'iire. 

Tlio  must  .siu'iiiflcaiit  fiwt  in  tin-  <-lcctfi)i'ar(li<tifrnm  is  therefore  that 
llii  htixr  is  Htiintirr  to  thr  upr.r  at  thf  hi  iiiniiini)  (liinirt)  and  mjaln  at 
tlif  mil  (T-inivr)  of  the  rrntrliulnr  ronlniition.  llow  mwy  tliis  be  ex- 
|il:iiii<'(l ?  Wlii'ii  ('li'i'trncardionr'niiis  aic  tiilu'ii  tliinimli  cleetnxlt's  i)lac<'il 
.lircctly  (III  tlic  Imse  and  a|)i'X  of  tlic  ventricle  of  tlie  exjiosed  heart,  it 
liiis  heen  foiiinl  tliat  the  eontoiir  of  the  eleetroeanlioirraiii  is  lii<c  tliat 
wiiieh  is  olitained  from  a  strip  of  mnsele  when  n  wave  (»f  eontrnetion 
I'lisseH  ah'ni;  it:  it  i-*  (ii|ph:i.sie  in  ehiiriicter  i  |iiiu:c'  ISSi.  a  result  which 
may  he  interpi'eted  as  indicating  that  the  Ai.ive  of  contraction  starts  at 
llin  hasp  ami  ends  at  the  apex.  This  rules  out  the  explaiuition,  at  one 
linn'  sujriiested  foi-  tiie  T-wave.  that  the  wave  starts  at  the  hase,  tlien 
|iroceeds  to  the  apt'X,  and  tiindly  ends  at  the  hase.  following  the  disposi- 
lion  of  tlie  muscular  fihers  of  the  ventricle  in  a  fohh'd  or  loop  foriri, 
with  tJie  111  nd  of  the  loo])  at  the  apex  an<l  tiie  free  eiuls  at  the  hase.  Al- 
'lioufth  the  exiilaruition  seemed  at  first  to  conform  with  the  einhryo- 
Iniric  fact  tlint  the  heart  is  developed  from  a  f  tuhe,  it  can  not  liold, 

;is  has  lieeii  shown  hy  ohservinj;  the  course  o  .e  excitntioii  wave  se- 
iiired  tliroujili  electrodes  placi'd  at  various  ))oinis  on  the  surface  of  the 
I  \ posed  ventricle  (pafie  1!(4). 

The  ex]>lanation  which  is  accepted  hy  the  imi.jority  of  observers  at  the 
[■resent  time  is  to  the  etfecf  tliat  Ihr  T-inirc  in  rniisid  Itij  thr  lonij.  ■■  ron- 
I'lHiancf.  of  the  tinlric  thamir  at  lh<  hnxi  of  the  rrntrirlr  than  <it  thf 
"li/.r.  To  test  this  hypothesis  the  criicial  experinu'nt  would  evidently 
I'c  to  sec  wliether  a  T-wave  eonhl  be  induced  in  an  elect I'oeardiotrrani. 
Mich  as  that  of  the  frojr  ventricle,  in  which  no  T-wave  exists,  by  hurry- 
in!,'  up  the  ••ontractioii  i)rocess  at  the  aptx  without  afl'ectini;  it  at  the 
li.'isc.  This  can  be  done  by  local  warminsr  of  the  a|)ex,  or  by  applying' 
'lie  ventricular  electrode  at  varyiiiir  parts  of  the  xcntriclc  in  an  excised 
licart  beatinir  in  Hins;er's  solution  of  relatively  hiiih  H-ion  coiicentra- 
'inii.  Mines  showed  that  under  these  conditions  a  typical  T-wave  ap- 
I'cars  in  the  elect rocardio^rram,  as  shown  in  Fisr.  S4.* 

The  existeiK'e  of  the  small  Q-wave,  indicatimr  that  the  contraction 
lues  Tiot  I'cally  start  from  tlie  base,  conforms  with  the  observaticni  that 
'lie  I'urkin.je  system  of  fibers  emls  about  the  i)ai>illary  muscles,  which 
'iii'icfore  wcuild  be  the  first   to  contract,  and   with   the  observations  of 

l.i'wis.  already  alluded  to  above,  on  the  appcaian f  the  iiejrative  vari- 

ilion  on  the  sni:'face  of  the  exposed  heart. 

The  mosi    important    cliniciil   ajiplicalioii   nf   ihe   dec!  rocardioiiram    is 


'This   Ir.'uinfcj    w.ts    fi-ntnl    ;r;i  mik   *Inn.r    left    !>>■    Vr<'U-^*"r    Mtnr-; 
rr!li^^in^    to    u>;r     it    thr    .mtlioi     is    in.Iri.t.  .1    (,,    th,-    .i>illi'.ritirs 


■  ■i    M.l.ill     rnin-rsily.    .-iiul     |gr 
'tf    iti.i!     iti*titnli.,n. 


i  ^1 


L'liJ 


fllK    niii  IHTIiiN    (IK   Till     lll<iol> 


:i-.- 


.(.     .Vu.uul 


II       Amx    ..,,!,, I 


'M 


,11 


(  .     Api A    waruitij 
fh.  ''.''■',\''!       '''"■"'"    ■''    <l"'r:'..i'.li-K.a.n    ,,„.|    ,u..v,,n,„l    „f    >.„tri,l,     „f    fr..^    ,l„.„iM>..    tlu.    »!. 


I  I.I  <  TIliH    MllillNiK  WIS 


•'ti.'. 


Illliliilllitfill.v    ill   rciMil)  I'lloll    with    Ihr    ili'li'l'initiatiiiii    iil'   Ih,    rnh    iif   timis 
iiilnsioil    iif   till    I  itildlinii    iiiiri     riiim    niilit'h'    tii    \ciit  lii-lf ;    llills,    the    I'  U 
iiiti'i'Mil,   iis   it    iiH   fjilh'il,    iiiiliriili'i    till'    tiiiii'    t.'ikfii    I'm-   thi'    iiii|MiUi>    t'l 
ii!i\«l  rroiii  the  viiupiiiiriiMiliir  tn  thi'  iiiiriiMih>\niti 'k'iiIiii'  iimlt'  iiiiil  liiiinlhv 

III  ili'lii.M'il   tniiiMiiissiiiM    this   iiiti'ivdl    I iiii's  nl>ii>ii'iniill,\    Iniiif.      Oli\  i 

Miisly  tilsii  <>iiiiitiliiiiis  III'  III  III  I  hliirl,,  iif  iiiirii  iiliir  jilinUnliiiii,  or  nl'  iiiirir 
I'lr  Ihilhr  will  III-  iiii.iiriliiili'ly  ri'vinlnl  In    tl lictiiiriiiirmLriniii.     'I'hr 

lllt('l|llftiitiiPII  iif  illilinl'liiHlitii's  ill  tlir  rnlilnlir  ill'  tin-  \  fill  IMiMlllir  purtjiiH 
III'  the  ('lir\r  is.  hiivM'MT,  not  mi  I'iisv  a  iiiiitlfr,  iiikI  slmitlil  iicmt  In- 
uinli'ltllki'll   Ulllrss  clllvts    I'lmn   tlli'   Unci'   Iciiijs  liiiM-  licrli   sciMlli'il,    fur   it 

Will    lie    riuiliil    tlijit     th rir>|niiii|iMi;    clri't  liu'ari|iiii;i  mils    ililTi-i'     rinill 

•  ■III'  iiliotlicr  ill  ilrtail;  I'lir  I'Xaiiipli'.  thi'  l{  uaxc  is  iisiiall.\  must  |ii'iiiiiiii)'iit 
ill  h'ail  "J.  altli(iii'_'li  siiiiH'tiiiiis  it  is  iiiiirc  iirniiiiiirnt  in  li'.nj  :!.  T  is  iilua.\s 
ii|iiii.'lit  ill  noi'iiial  iiiili\  iiliiais  in  cuim'  taki'ii  liniii  hail  •_'.  lint  it  is  imt 
inrrcillli'ii'ly  iiiM'ittil  ill  tli.is.'  Ill'  h'ail  :!,  ami  may  slmu  partial  iii\  iisimi 
III  tliiis<'  .  .a  h'iiil  I.  Till'  (^  US  k'liilip  is  iil'tcii  lit'  pci'iiliar  I'lHitnur  in 
iiirvcs  fi'i  .  I  h'iiil  ;!.  'i'lii'sc  \aiialiiiiis  arc  piissilily  ilipfinli'iit  iipnn  llir 
Illative  prcpnnilt'iai..  •'  nl'  tin'  miisciilatiiii'  in  the  Icii  ainl  li^lil  vcn- 
iiii'h's,  fur  it  is  cviili'iit  ihal  ihi'  anniunt  nf  iniisrh'  ini-liiih'il  in  tlir  path 
\»ay   iictwccn   tin'   t\M>   Irails  uill    \ary. 


CIIAI'TllK   XXX 


■I  I 


(I.IXK'AL  AI'I'I.ICATIOXS  OF  rHIJTAIN  l'I[YSU»LO(iI(; 
.MKTIIODS  (CuutMj 

CLINICAL  APPLICATIONS  OP  ELECTROCARDIOGRAPHY 

The  Electrocardiogram  in  the  More  Usual  Forms  of  Cardiac 
Irregularities 

r.V  K.   \V.  SrolT 

Tlip   prinriplc   of  tli,.   ii|.plir,itii.ii   „{   tlic   >tiiMi:   ■r;ilv;iiii)iiictfr   Ic   tin 
siti.ly  .,f  ciinliar  iririrulMritics  Iijis  Ih-cii  iiidicMt.'d,      It   is  our  ol.ject  li.M- 

to  outline  some   of  tli,.   mon imo,,    f,,,.,,,^  ,,|'  inviriil.-ir   iifai-f   ix'tion 

uitii   a  l.ricf  (I.M'iiptioii   of  tli.-  almoniialiti.vs   in   the  (■{.■.■troi-anliosrinm 

ivsiiltiriir  tiKT.tVoiM.     l-"or  tli.'  saisc  of  c pai-isoii  a   iioniial   <>lf.-tro,-ar 

(lioirraiM  is  shown  in  Kiu-.  s-_'.     Tlic  raiis,.  an.l  ivlationsliip  of  tli.'  various 
<li'llrcti(.tis  lia\i'  Im'cii  cxplaini'd   (  src  pairi'  iMi^). 

Sinus  Arrhythmia.    This  inviruhnity  is  s,.,.n   c-oiiiiuouly   iu   fhihhvn 
an.l  youn^'  adults,  an.l  is  uitiiout    pathoh.-i.-  si-nilican.-o.     'I'JU'  rlfcti-o 
canlio-rrani    j.roscnS   tlic  normal    di'llcctions   and    sliows   l.y    the    varyin- 
spac-s  iM'tucrn  the  I'  d.'llcc-tions  that  th.-  eardiar  iMipuls,.'lias  Kcmi  --•u 
cratfd  at  sliLrlitly  irri"_'ular  intervals. 

Sinus  Bradycardia.     The  eleeiroeardio..nani  in  a  simple  ease  of  simis 
ln-ad\ear.lia    is  usually    normal,   exeept    that    the   delleetioiis   ...vur  at   an 
IMMIsually   slow    .ate   i  Fii:.   S.",,.      This    iudieates   that    the  .Mrdi.ae    impulse 
is  built  up  at  a  sh,u    rate,  Init   uher   i.',iierated  it   evokes  a   normal  aurie 
ulai'  and   M'ntrieidar  eontraetion. 

The  Extrasystole.     The  extrasystole  may  I it  her  aiirieular  or  ven 

tii'''ilar  in  oriuin.  Meeasi,,nally  a  rare  type  is  seen  iu  whieh  the  iin 
l.uls,.  arises  iu  the  jnuetional  tissu.'s  hetweeu  the  auri.de  ami  veutrich- 
When  til.'  f.ieus  ,d'  impuls,'  iirodueti.in  is  at  ..r  near  the  sliio,n(rir„hir 
iio<lr.  the  tvsultin..'  .•l..-tr..e;,r.li..^Tam  .•.uMpl.'X.'s  ar.'  praeth-aliy  n.irmal 
If,  hou.'ver.  the  seat  of  impulse  lormatiiui  is  rem.ive.l  from  the  .S  A 
H. .,].■,  th.'  I'  .lelli'.-ti..u  may  he  .listorted  ..r  aetually  inv.'rte.l.  f.dl.iw.'d 
iiy  a  n.irmal  (^  HS  T  .'.imi.l.'x  i  Fii:.  f^ti  i. 


!i!  t! 


le  ease   o 


in   either  the   litrht    or  the   left    veiitt 


"•   »  •• 'J^^^;/.^/^;I^  ,    liie    ejirdiae    impui 


>!■  orimnali 


This  aliuurma!   site,   to._'etl; 


n.INirAF,    A1'I"I-I('AT      S"S   OF    i:i,i:cTKO(AKIiI()(lHAPlI Y 


2H7 


t 


;LA. 


Si  mis    ljraiiyi.ar<iia.      Kate    .'J 


ir    tMiiri 


cat  ilioyi.Tm.      I'  K    iiiitt\;il     -.17 


t«.       N-'tc    the    imrMial    ai'pr.iraiicf    of    ihi?    ilcitr'>- 


Sr  -R 


~^ 


.Xw 


T  f 


r--" 


I 


A'lt'.Ml.ii     .  •.  tt,i-~\ -.ti.t.  ,       'l\r,,    ;..iiifiil.ii     )  v.tia-.wt..!.  >     t-'H-v.  lilt;    Iv^m     ■■.mhi.iI     < ''mpl' x*'*- 
Ni'tr    ill.-    .Mii|.i.     -MU-TM    nt    liM     .  \tiasysti.l-    ii-'Iiiar.  ■!    i-^     Iii>-    in\t--;-.n    ,.t     1'. 


pig.    87.— \'cTitrit.*nlar    c\iras>vt..|i  s    aii'-iiin    in    \\:r    rivilit    \iiitritli- 


l-lj^.     S."-.       WlltriLllI.u     r\tl  .l''>  »t.'U-     .11  iMiiH     U.     till-     Ictt     \rlitl  U  1(  . 


.j<r.  ■ 


iiJiA/*^  ,v 


If  u 


21  IS 


Tin:  ciiicrr.ATKiN  op  tiii:  r.i.nuh 


witli  111.-  p.'itli  uhicli  til,.  iiii|iulsc.  Inkcs.  |>i'<ii|i s  :i  nnu-li  ^r||.;it,.r  diiV,., 

'•' I'  '■■'■'■trie  i.,,l(iili;il   lliiiri   is  seen   in   lli,.   noriii.'il   clcctrociinlioi.'rjin 

Wluii   the  iiMpiils..  aiiNcs  ill   the  lit-lit    \  ciiliiclf  lu-ar  (lie  l>ilso,  tlic   |>ii:i 


\i   :i 


i         } 


*?        R 


^it-QMM^W^ 


^s/SM^/m 


T  ZiT  i    -ell 

p  .    r 


i    k 


l'.^.    .^'i.      r.u-Ms.vM   ..I    ia.h>,^,..!i.,,       An.i.nl.n    ..,wh,        N.„,-    M,,,l    lli.     /     ,!,H,,i,.„;    Till-    !,,„  k    ,„.     / 

k.iU-    Jiin    JHT     intMiilf, 

cipal  I.'  (Icll.'ctiuii  is  iipwaiils  in  Imth  l(.ai|>  1  aii<l  2.  Aiisiiii;  ni'ar  tl" 
a|ic\,  thr  prill. -ipal  I!  (Icllcrtiuii  is  up  in  Iracl  I  ami  di.uii  in  lead  12.  Tw - 
cxtl-asystdifs   lioth   arising    in    tlic   rivlit    Vfiitiicic   arc   slmun    in    Ki...   s; 


I  •«  ■ii'*i|«W|^iS«i*w<»i 


I 


R 

I 


I-.  ' 
'■  I 


,^  Vij.  'M.  ^A,,ri,ul.,r  M.nll.ui..,,.  T...,.!,  1.  .■,  ,1,  N,,,,  ,l„.  ,,,,„„.  |,l„,n,,;i,„,  „.„.,,  i„,,,,,„  . 
,',■,,',;';',;,";„; '"'     •'-""■    ■"    •"'>    '■    .I.H..tl-M.-    ,„    ,-.  I..t,.,n    ,,,    ;,-.       AK..    U,,     mi,  .,,k,I    -y.,.:,,^    ,.,    ,1:, 

In  tlic  case  „(  III,.  |,.tt  \,.ntiiclc.  a  l.asal  impulse  yivcs  a  dnwnwar. 
principal  .Icll.cti^^n  in  !,-a,i  !  an,l  up  in  i,,..i  _'.  Wiicn  li„.  aiMTrant  f. 
PUS  is  located  near  tlie  apex  oT  tlic  l,.|t   ventii,-!,..  iji,.   principal   dell,., 


CI.IMCAr.    AI'I'I,1(ATI()\S    OF    l:r.i;(Tli(l(AI{|)IO(lUAl'EI Y 


'2fi9 


ticiii  is  (Iciwii  ill  lidtli  Icids  I  Mild  '2.  Any  (iiit'  of  scvciiil  of  tlio  irciici'iil 
ly|>cs  of  cxt  |-;|syst(ili'  iiuiy  occur  in  tlii'  siillic  ]>;iticii*.  I''ii.'.  SS  sliows 
MM   cxtr.'isystnlc   orii;iiiiilimr   t'rnni   tlic   left    vciilficlf. 

Paroxysmal  Tachycardia.  Klcctroc;ii<lio<:riiiiliic  i-cconls  tiiUon  in  tlic 
nitorvnl  liotwccii  llio  jiiii-oxvsiiis  iiniy  iippciir  iioimiil.  Diiriiifj  llio  tadiy- 
iiu'dia  tJio  rocords  nni'ni;illy  sliow  only  two  deflections,  R  ;iiid  a  rnmliinn- 
linii  of  T  and  tlio  siiccccdinu'  V  (I''!-,'.  S!)i.  If  tlic  i)iirox.\  sin  is  of  iinric- 
iilar  (M'isiin,  tin-  I*  dcllcction  in.iy  lie  inverted.  indieMtintr  tlint  tlie  new 
inciis  of  iiiipnlsc  ]nndnctioii  is  lociited  at  some  otlier  site  tlian  the  sino- 
amicnliir  node.  Karely  tlie  new  focus  may  l)e  in  tlie  \-eiitrieles.  Heeords 
;.iken  diirintr  the  i)ai'oxysiii  may  siiow  a  rapid  suecession  of  dellections. 
■-iiniil.iliiiLt  isolated    veiil  rieular  extrasystoles. 

Auricular  Fibrillation.  Tlie  elect rocardiouram  in  aiirienlar  (ilnilla- 
iiiiii  shows  thi'ee  dislincti'.e   features: 

1.   .Misence  of  the  1'  detlectioiis  txpical  of  auricular  eontraetioiis. 

'J.  The  \ciit rieular  complexes  i<^-!i-S-T  \\av<'s^  occur  in  irreirular  so- 
'    Hiice  ami  may  \ary  in  heiL'ht. 

I    The   |)reseiiee   of  siinill    irreirulai'   oscillati'  hest   seen   lietwoeil   the 

'iilricular  ('oiii]>lexes.      .\   tyjiieal   traeintr  of  this  eoinlitioii   is  shown  in 
l"i-.  !•(>. 

The  de|ien<leiic(>  of  the    I'  \\a\(>   u|)on   auriiMilar  eontraetion   has  lieen 

iidic.-itcd    (  pa  ire  2()1).      Its  ahsi'iice   in    anrieul;ir    (iliriliation    is  accounted 

'■■'f  hy  till'  fact  that  liie  iiiilividiial  muscle  (ihers  of  the  auricles  eonti-act 

i':i|ependentl\-  of  one  another,  so  that  siune  liliers  ai'c  in  a  state  of  con- 

'  :ii|inn  while  othi'i's  are  relaxed.     This  renders  impossilile  a  coordinato 

■  "iitraetion  of  tlie  .•niricle  ;is  ;i   whole. 

The  multiple  impulses   from  the  lilirillalin;r  anrieles  reach   tlie  veiilri- 

■1";  and  evoke  a  eontraetion  provided  tlie  ventricle  is  not  already  in  a 

'.itc    ..f    colli  r.-ictiiiii     I  refrai-tnry     peric  d.     paiii'     l..-ii.     These     iircrnlar 

utricular    responses    will    of    course    )»r(Mluce    unei|ual    s|iaciiii;'    of    the 

•utricular  complexes   in   the  clei-trocardio^ram.      The   xariations  in    the 

"i'.'lit  of  the  I{  dellei'tioiis  is  tlionuht  to  he  ilue  to  the  distortion  ciiiscd 

the  sui)erimi>osition    of   the  small    waves   representiii'j:   auricular  ae- 

■  -ity.      These   sni.'ill    waves   must    ocelli'   throiiirlioiit    the    whole   ('ardia<' 
'■le.  hut  are  more  or  less  masked  liy  the  \-eiit  ricnlar  comiilexes,  appear- 

i'-T  as  separate  oscillations  only  diirintr  <liastoli'. 

Auricular  Flutter,     .\nrieular   llntter  was   discovered   hy   the  electro- 
'dioiiraph.  and  it  is  practicallx    impossilile  to  make  a  diaeiiosis  of  this 

•iidition   without    the   use   of  the   strinir   uralvaiioiiietcr.      The   auricular 

i'lleclioiis  are  usually    rhvlhmic  and    in    the   avi'rafe   case    \ary    in    rati' 
'iiii  '_>(l()  to  :'.."((  per  iiiinilte.      'i"he   initial    dellection    of    I*   m.-iy   he   li.ise 

'■uati\e  or  apex   lieu'ative      lip  or  down     dependinjr  on   the  site   of  the 


imuaM^ 


!     1 


Tin;  (lunr. Alios-  of  tiii.  iii.oon 


■I 


i>rij?in  (if  the  ;iiiiiciil;if  iiii]>iilM'  i  « lioii  .-n-isinu'  iVdiii  some  ntlier  suuii . 
than  Ilic  S-A  iioil.'  till'  iiiipiilM'  is  Slid  to  lie  fctdpic  >.  I'sudlly  a  rosjul.ii- 
supocssiiin  (if  1'  (Icllcciidiis  can  ho  traced  thnuiirhout  the  record  iFi.' 
ML. 

Since  it  is  iiiipd.ssililc  for  the  ventricle  to  res|t(iii(l  to  all  the  impiilsev 

coiiiinfr    fidiii    tlic   aiii'iclcs,    a    iditidii    of   partial    lieart-bloek    ohtain- 

(2:1-  :i:l  4:1,  etc.'.  The  \ cut riciilar  coiniiicxes  will  occur  roffiilarl) 
except    when   a   :!:■_'   rlntlim   exists. 


'■"-■     '''        ■^'"<    I'l-"    ll-.i'-lT       .\Mri.uI,,r    Tal.     tn.l       \",il!  i,'i;:.,r    i.d,-    Sc        \,ii,-   III.     inv.-.-i.Mi   of    ill, 

ill  11.1  ll, .11^. 

I'sually  the  \ cntricnhir  coiiiplexes  are  such  as  to  indicate  that  tli 
stiiiinliis  arose  in  the  auricle  ( supraventricular^  The  hcitjht  of  tip 
individual  detlections  (^-f{-S-T  may  vary,  depeiidiiiir  on  the  predoniinani' 
of  a  rii:ht  or  left  ventricular  hypeitroi>liy. 


ir-fi  ( 


;i  II 


'■'«     ''-'■      I'.uH"!    >'  I    ;.i.n,.ii,       N.,1,-    111,-    ;,...•■,„, I    ai,],.  ,,.,,,,,,     ,,f    i|,,.    ,-l,-,  li .,.  .ii  ,Ii.iki.iim    ,-v,.>|.1    l 
ih.'    |,i.,I...n::ili..n    i.l    lli.-    I'h'    iiit.Ti.il.    uM.li    iiM-.isiir.  -    .:?,!    ^^cininls. 

Heart-block.  Tluic  ai'e  three  dcorees  of  se\crit\-  in  hearthle  k:  i  I 
delayed  ciinduetion,  ili  i  partial  dissociation,  and  i :!  i  coini)lete  dissoci;i 
lion. 

Any  one  of  thesi nditioiis  nia.\-   he  jn-espnt   in  the  same  patient  ;i; 

successive  intervals. 

l)i;i..\vi;i>  CoMiiCTiON.-    When  the  iductiii!,'  tissues  of  the  heart  ar. 

so  afTecte(l  as  to  cause  an  ahnormal  i>rolon>_'ation  of  tlie  I'-H  inteival 
the  condition  is  called  dela.\ c-l  ciniduction.  The  ventricles  respond  t' 
each  sliiinilus  oiiL'inatiny  at  the  sinus  node,  hut  the  time  re(|uired  for  th' 
impulse  to  |)ass  tlirouu'h  the  eondiietinir  tissues  is  loiiijer  than   norm:: 


CI.IMl'AI,    M'l'I.ICMiliNS   or   KI.I'CTKOCVUDIOC.KM'IIV 


271 


la  a  siniiilf  case  flio  ('Icfti-dCiirdiofirain  may  appmi-  peifcctly  normal, 
l>iit  when  tlic  riv  interval  is  incasiircd  accurately,  it  will  be  found  to  be 
lengtlieiii'd  licyond  tlie  extreme  limits  ol'  the  iKirmal  i0.20  seconds)  (Fiff. 
02  I. 

Paktiai,  Dissoi  iatkin.  ^In  tlic  typii-al  case  of  paitial  dissociation  the 


X         ' 

:         :   '..  :       ■;■      "-;:   ■ 

I'H.    '»3.      I'artill    ilUsoi-i.itii.n,       N..ti'    t!n-    f;ii!iiri-    "f    \>r;triin!.r     r«-|.Mii-,-     I'mIIcwhiw'    ttif    -rci-li'l     /', 
which    has    lii.ii    i-ix.-.U-.l    hv    tw.i    i-itia-) -t..li-i    <\i    ..t    v.i.ti  i.  ular    ..riuiii. 

Nciitrielcs  I'esixnul  to  the  impulse  comiiii,'  from  the  auricle  nii'^l  of  the 
lime,  iiiit  occasionally  fail  to  do  so.  when  the  condition  is  calleil  ■■droi)ped 
Ipcat."  The  elect rocai(liov;ram  records  a  1'  dcllcction  but  no  ventricular 
complex,  slio\viii<r  that  tiie  auricles  have  contracted  at  their  usual  rate 
init  that  the  ventricles  failed  to  rcspmid  to  the  stiuuilus  coiuinjj  frinn 
the  sinoauricular  node  I  Fiir.  !•:>>. 


;^'    'M.-  l'Mi;;j.li-tt'    ilis-ioi  iatiim.      Niilf   that    tilt*    /'    u.i\f    >iatfs    iiLi.ilarly    aii-l    licar  -    ni   th-liiutc 
Intiuii   to  the   A'   «avi-   ■■{  thf  vriitriiiilar  ..imi'hv.      .\iiikii1:i:    r.i'.f    TJ.      VVnlrKiihlr   rate   40. 


CoMi'i.KTK  Dissociation. — In   a   simple   ease   of   complete   dissociation 

lie  auricles  beat  imleiicndently  of  the  ventricles;  hence  the  P  dctlection 

if  the  elect rocardiofii'i"">'  bears  no  relation  to  the  ventricular  complex 

<i-K-S-T)   (Fi^'.  !Ui.     The  1'  dcllectio:is  s|)acc  resulariy  and  are  easily 

made  out  when  thcv  fall  during'  diastole  of  the  ventricl.'.     Occasionally 


M' 


272 


Tin:  ciun  i.\Ti((\  ok  tmi;  nLoni) 


tlic  ailiii-l..  will  |iM|.|.,.|i  t„  cniitiiirt  .liiiiii-  v.'iitri.-iil;ir  systclc.  ciillsins;  ; 
(listc.itic.ii    of   ll„.    vciit.iculiir   .•.,ni|.l..x    \,y   tin-   siiixMiinpositioii    ,>(  >i    ]■ 
'l<>l!<'--ti.m.     Hx,.,.,,t  Ml„.n  this  (..•.•iirs  tli,.  (^|{.S-T  c.nii.U.x  is  flio  ncnrin^ 
sM|.i;iv,iitiicuiar    t.\  p,..      Th.-    1>   .l.llcctiolis    ocrnr    llioiv    fivr|llcnlly   tl,;,i 
tile  (^K'-S-T  n.iiii.ifx,  sliouiiiif  Hint   the  fiiiiicl.'s  iiif  l.catiiin  iiioiv  oft.i, 


tl.iii  the  \cntriclfs.     Tiic  aiiriiMiliir  rate  in  1 


iif  avciairc  case  of  CDinplc 


hcart-l.l,„.k   is  .•il.niii   72.   nliilr  the  \  ..utricular  lalf   is  iin„-li  slower   cr, 
lo   JO). 


P     ' 


1|^  i) 


J 


:l        ij      H 


CIIAI'TKI!  XXXI 

riJXKAL  AI'I'I.ICATIONS  (»l'  CKUTAIX  I'll  YSI(>l,0(iI(' 
.MKTII(»I)S  (Coiif.!) 

POLYSPHYGMOORAMS 

In  t.-ikiiijr  pfilyspliyiriiioiiiiiiiK.  tlic  I'dlli  w  iiiir  tccliiiic  is  usuiilly  rnllinvcd: 
Venous  Pulse  Tracings.  Direct  the  (pI)m'ivc(I  ikisou  tn  lie  down  witli 
his  lic.-nl  sliiilitly  i-iiscil  liy  a  cnsliinn  iind  liciit  tn  tlii'  riiilit  side.  I'laco 
tlu>  receiver  (lliistie  funnel  i  over  tlie  jnuMi'iir  hull)  en  lln-  linlit  side  nf 
liio  neck.  This  lies  iininedintely  ali'ive  the  inner  end  ul'  llie  clavicle. 
I'lrini;  the  style  of  the  lecordin'r  tandiDur  to  \\?it<"  with  a  minimal 
amount  ol'  friction  on  the  paper  or  drum.  Since  a  venous  pulsi>  traciiiir 
can  not  lie  interpreted  uitliout  a  simultaneous  traciusr  from  an  artery, 
now  adjust  the  liutton  (d'  a  receivinu'  tamhour  over  the  radial  artery  and 
adjust  the  style  of  its  recordin;.'  tandiour  so  as  to  write  on  the  drum  ii' 
tlu-  same  pcri)ondicular  as  the  style  of  the  venous  tamhour. 

T-aciiiTs'  should  he  taken  with  the  recordinu'  surface  at  a  moderate 
sped.  P>efore  disturhiii<,'  the  relativt>  positions  of  the  writinir  points, 
allow  thom  to  make  vertical  marks  (with  recndini:  s\irface  stationary  > 
at  various  parts  of  the  tracings.  These  ali'-'ument  marks  ])erinit  of  ac- 
curate coiTiparisons  lietwecn  the  cuives.  Hepeat  the  ahove,  rising'  the 
carotid  instead  of  the  radial.  A  time  tracin-i  ( ' :,  >*i'<'- '  shoidd  always  he 
taken  simnltaiK'ously.     The  i)olysphyenio<iiiiph   is  shown   in   l'"iir.  !»."). 

To  iiderpret  the  venous  curve,  make  a  vertical  mark  on  the  arterial 
pulse  tracin;.'  corrcspoiidinir  to  the  heirinninu'  of  the  jinlse  upstroke.  II 
this  is  d(UU'  on  the  ratlial  pidse  tracing:,  nu'asnrc  one-tenth  of  a  second  in 
front  (d'  it.  and  make  a  vertical  nmrk  to  allow  for  the  time  lost  in  jiropa- 
-atiou  of  the  pulse  from  the  heart  to  the  radial  artery. 

This  line  :*>  (corrected  in  ease  of  radial  i)ulse^  cirrespouds  to  the  he- 
'.'iuninu-  of  the  s|ihyt;mic  i)criod  of  veutricul.ii'  systole-  i.e.,  to  the  ojk'u- 
iiis;  of  the  somiluoiir  valves.  Measure  the  distaiu-e  fnun  it  to  the  near- 
est vertical  liiu'  that  was  made  to  iuilicate  the  relative  jiosition  of  the 
writiiifr  points.  Then  measure  ofl'  the  same  ilistance  in  the  venous  trae- 
iii-,'  from  the  correspondiu!;  iudicatni-  line.  This  v.il!  fall  at  the  !>c<rin 
ninir  of  the  small  wave  ((•»,  which  is  due  to  the  liulLrinji'  into  ihi-  auricles 
of  the  closed  aurieuloventricular  valves,     ^\•'\<z.  'Ml) 

27S 


m 


n 


«  '■ 


i: 


I 


:^i 


s  1 


Tlif  a\ii'ii>ulnr  wav 


TIIK    CIKCI  I.AIhlN    (IK    Tin:    IILIHID 


0  ((/I  occurs  oiicliflli  ,,f  a  sccoml  in  lYoiif  of  c,  a 

aiico   ill    front   of 


may   now   In-  asctrfaincd    hy   iii,a>uriiiir  otT   this  dist 
Tliis  is  line  1. 

Now  mcasuiT  the  distai n   the  radial   pnlsc  tracinjr  fr.mi   the  h, 

ninir  of  the  n|istroi<e  to  the  .licrotie  notch.     The  distance  l.etween  tl 
the  sphyt'inic  period  i  !• 


lese   jv 


(■■(•.r    lr...iMK«.       Tlw    ,:..,;,«.,, k    ..|.,rii,,     „     .  „ ,  .1,1..    \        ,"■""""'".  -""I    P<rni.iMi-iit    ih.m    M„„k, 
r.."„|,    ,„    ,|,tT.r,-,|.    ,,,M.U.  ..1.1,1,1,     .|„,,N.    ,„.rinMI,i,K    tin-    l:,kir,«    „f    ,.r„trn.l. 


^   -Measnrc   ofl'  ,h,.  same  distance   on   the   venous  tracint;  fro,,,   r      hinr 
•'  Will  he  lonnd  to  fall  just  hcfon-  a  small  wave  (ci.  which  is  due  to  the 

'!"'•';:"  ' ""'-  "'■  "'-  "•'•••'-I'i'l  valves.     This  praeticallv  coincides  ui,l, 

the  dicrotic  notch  on  the  radial  pulse  tracinfr.     Sometimes  a  little  uav 


i'tK      ■'(►-  -\„ 


I  I  r-in:     I,.     I'.     ^  artcr   i 

omns  on  the  upstroke  of  .avc  ,•  just  .here  line  ',  falls.  This  -o 
n  .•..ies  with  the  closure  of  the  semilunar  valves.  The  distance  I.etwoen  i, 
ana  wave  r  .-orrcsponds  to  tie  postsphyKmi,.  period 

Ihe  cause  for  the  depression   ,„„,,!,„,  ,,    f.,„„,.i„^  ,,   ^^  ;„  _  ,^^, 

'l..st.H.l  hy  reterriii«  to  the  intraauricular  curve   ( !■%.  97).  ,o  which 

s  .liready  CN,,lained.  the  venous  pulse  tracins,  is  .p.alitativelv  similar' 


un 
n 


r(i|.YSI'IIV(iM<>(lH\\IS 


riir  rise  in  llio  I'liivr  t'olldw  iii(j  (Icpifssinn    ''  i*  «' 
ilif  iiiirifl*'  with  lilood.     'riii>  u"''^ 
^M  II,  nlliiwiiiy;  tiic  MimmI  tn  f.ill  into  tlif  vciitriflf. 


27» 

iUlscil   liy   tin-  lilliiiH  "t" 
until  r,  whfii  tli<^  tricuspid  v«lvc^ 


l»I      'I/-.  -  knlun.l     ll.i.uiB^     (l .,.l..li.i.     .r.n.i.     Miiln.l..     .iiiii.lr     .i;.l     jtirfMl.li.     t..     stliiw     the- 

K,i„i..l    rrlatinn»lii|is   ..l    Ih,-    sat -    «:ni-.      An    ,!>,li..,.ii.li"Ui.i"i    i-    :.l-.    -h.iuii.       N'.v.'    llwl    Ihf 

iit|4ulai    anil   atiriiillar    iutvi>    ha..'    Ili'     -.tin.-   t..nt.. 
iliiiiiiii    ..y«t"li-    "I    llu-    vpiilM.li".        lAlt.r     l.t«i»   I 


1.1    tlial    111.     .l,|.it-»tM 


in    til.  Ill    "t  t  III.. 


Til  iiitcrprct  tiic  caidiojiriiin,  :i(l.ju>.t  ii i\iiijf  tMiiiliouis  to  lln'  I'lnlial  ami 

npcii  lu-at.  witli  both  writiiii;  stvlcs  in  llif  saim-  piTpcmlicular.  ami  f(illi<\viii>; 
tiic  otlitT  (lircrtidiis  iit'sciilM<l  iiinii'i-  "xenons  pulse"  iiiaik  on  tiic 
earilio^niiii:     (See  Fifj-  I'f^- ' 


m 


rm-i 


.1  m 


■  ^  ..'vv- 


fVJ 


\  J 


■^'r -'-'■' ff'^-'.i 


J  J 


_,-_.-  :;-^ 


!:s    ■JX— l'.,lw|.|.y^ii..,^i»,.-,i   i,,,|.i.|ii,|{   ].ijc.i!..r,   .-.prx   av..l    r.i:!ia!   tr.-.tini:-!.      !."!••    4   'V<    tl..-    i.i.lial 

•iJung  iK  tirst  of  ail  Icicatrcl.      Il   i^  tlit-n  lr:iTi«ffrriil    lliy   im-asuriMiii-iit    fr.im   tlir  alimimi-iit   mark   on 

-     rii[ht   oilnf   of   the    Iraiinui    tu    t!ic   jiiuular   aii.l    l/in    sfoiii.l    siihlractt-.l    fr.im    it.    KiviiiR    line    .'. 

■.\  i.n   till..   IS  siinii.irly   traii-f.rri-.l   to   tin-   .iprv    traiiiij!.    it    falls   s"mr«lnT<.   ,im    tin-   npilii.Le   llu-    I'r 

^ iiig  iif   wliifli   is   line   J. 


H 


;l 


I  I  ji 


f«  \i 


M 


L'Tti 


Till     (  IIJll   I    \TlciS    (iK    •nil      Ml, Kill 


I.    TIm'   hriiimiil,.,'   nf  III,.   .|,|ivi,|,|i,.    ,„.,;,,, I    ,|.;,    ,,|,„,   -  , 
■-'.    Tlir   rli,l    ,,f   ill,.   ^|,|l^.J|,|i,■    |.,iin,|    i  K  ,    ,|j||,.   f,  , 

•!.     'I'Ih'     .•ll|li|.|||,||.     w  ;|\  (.. 

».    Tlir    lM..,ini,li,..^   „!•   v..|,t,.i..„l,.,|.   svsl,,!,.    Mli|r,.r,.,„.,.    Ih.(„,...„    I    1    I 

'■'|ll;iU    |i|rs|,||\  Mini,.    il,),.|\..||' 

•'■   '''' l-"i'"-'"f.inii.M,|.,\,.,it,i,.|,i..,i-  M,!v,.s  .|u^^,.>t  |„,i,„  ill  |,.;i,.ii„y 

•^''"■'^  ''''^ '  ""■  "■■"•'"-  ""■  •■^•■"•'   '" nt  ;,I   ul,i,.l,  ,|,„  I,..,,,  ,.„„„|. 

.■Ill'   li<;ir,|. 

"   '^  i""""'^""  <"  'M-ik,.  ..,.,,i,ii,   ,|,;,t   III,.  l.iitt.Mi  .,(•  tl,..  t.-nnlM.i,,.  U  ;„• 

'■'"■■■"'■'"    '•"■''    ""■    ■■''"■^    '"■■•".    -i" llHTuis..    ..|    ,|,.p|.,..snl    nr    ,|,■u'ilti^. 

".•IV..    lii;iy    l„.    iiiscnl,,.,!    Ill    vriifii,.||l,.||.    ^\.,|,,|,. 

Simultancnus  Arterial  Pulse  Tmcings!    Tl,<.  vrLu-itv  „f  tl.o  trmsmi, 

' ;"' "'"  '"''^''  "•■'^"  '^  '■■■'I'-Hi.-ii-.i  i.y  i,i,...Ki,i.Mi,,,  ,1,.:  ,i,„„  i„,,„,,„„  ,,„. 

'•"'"'"■    "-    '"    ""■    '•^'"■'i'l    ■■"■'1    i"     111"    r.l.liMl    ;,,l..,i....    tM„.iML'S    u(    „|,i,.„ 

'"'"    '•■''^'■"    ''■'■    ••'IM'l^iM'-'    nt„.    r |\i|,..,    t,||„l„,„,    ,,,    ,]„.   ,.,.,,,,,i,i    .,,,„,,.    .,,,,1 

.■Mlnllli   r    In    till,    llhliiil    ;|||,.|.v. 


I      ''I 


k  :\ 


It^; 


Abnormal  Pulses 

''"'"■  l'"ll"^^ii"-'  i-.'i  liiii'r.li-M.|i|,ii fi|„.  i,i,-,in  i'l,;ir.-i.-t..r.,f  ..iLiMn,,.,! 

I)ilki.s: 

The  Ventricular  Pulse.     In   tins  „„  •'„•'   „„v,.s  ,yr   pivsi-nt   in   tl„. 
,ni'-'iil,.|r  tiiinnu.,  tlir  Inint   .-iHi,,,,  l.-im.-  ,.ii|„.|.  i-,...,nl,„.  „r  im.-i.liir      I„ 

;.'".  '"'""■'■  "■•'"••  ""•  •''-'■' '■  •!"■  '■••'■•  "-nrs  iiiiiy  ,1,.,,„„,1  „„..  1,^  „,,.,. 

illni^uMl,,.  nd.tiHiri,.!,..  ,2  iiMM-..;,..  in  ,|„.  |„„,,  ,.,...„,,:!,  „..„„,,,„„ 
li".-nt-l,l,H.k  .■,ss,„.|.it,.,|  uiili  .■|i||.i,.|||,„.  Iil„ill;,tinii.  \Vli,.,|  |I„.  li,.,.,it  i- 
.m...nN,|..  ,1„.  ..|l,s,.|„.,.  ,„•  ,)„,  ..,-   „,.,^„,  ^i^„i„,,^  iiiiri,.,,),,   (i|,,illMtii.n 

Delayed  Conduction  and  Heart-block.    Tl,is  ,...,„s,.s  m   ,.),,., i„  ,|„. 

','""■  •■'•'■■" ^''''l'  "'■  ""•  ""■■  ^""1  ••-••  ".M.s  in  ,|„.  J,|.|,l,.|,  ,1,,,;.     xv|„.„ 

;'"■  "'■■^"■"''"'•><  i^  "'•  'iH'  first  .l,...a,.,..  ,!„.  ••,■1.,.-  iiif,.n,i|  n,..r,.lv  i.-.i-.n.. 

l-.iml,rni.,l,  l,nt  ulu.i,  ii   i.  „fsi„.|i  .!,.,,,.,.  ,|,„  ,i,„  „„,„,,,  i,„,,;,,^„  ^ 

tiiMis  tiiiU  to  1„.  ..imvi-^..,!  .-ihin^.  ,|„,  „nii,.„i„v,.||||.i,.„|,,,  iMnnjI..  i.i.hitnl 
■■t  w.v,.>  ,.|,n  I,..  ,|,.tiHi.||.  in  th,.  iii-.M,,.,-  ,l...,.|...,.s  „r  Ii..,.||.t-I,i,„.k  tliriv 
••nr  iT.'nl;i,-ly  ri'i-nnii,..  "..|-  ,,,v,.s  i,,vii|.,,  |,„  ,.„|,,,Mnt  liin..  ii.|,-iii,„Kl,i„ 
>"  II"'  ••«■  Nv.'ivis.  ]■;„■  th,.  |,ui|M,s,.  „r  ,.x,.„.|  .-Mnily.sis  i,|'  tlio  n.rvi.s  In 
.s,is|,,.,.,..,|  ,.,.,..  ,„■  ,|„|,,„,|  ,.,,„|„,,i,,„^  I,  j^  ,,,.,^,^^  .•„|v,-iiit.-,o.,.„i,s  tu  ,ln,  , 
viTt,..;,!  hiM.s  l„.l..u-  .1,,.  tr,-,..ii,.,-  |.,.,.,.,.s,„,ii„_r  ,1„.  |„...i„|,i,„_,  „,■  ,„,ii.u!,-,r 
:"■■  ;''"l'-'.M.lar  M.tnl,..  Tl,is  Inis  I,,.,.,,  ,i„ni.  in  tli,.  (m.-i,,.  ,..,„.,„li„.,„| 
III  Fur.  !)!).  ' 

TIh.    Ii,„.    .i„iiiin-    tli,.s,.    two    M.,ti,.;ils    in,li,;it,.s    fl„.    , h,,..;,,,,    ti 

..>•■•.■,..■•    inti.iv,-il.      Wli.n    it    |.M.,.,.,U    „i„.  liftii    „,•   ,    „„..„„,^    ,,„.,..    j^ 
<i<'lay  111  ill,.  I'oniJiii-tiDii  liiiii'. 


Inl.V^niMIMiXiU  Wl.s 


.'77 


A  li'in'iiii.'  sliiiw  iiii.'  .1  liii.'l.i'i'  (li  ijii'i'  lit   liriiil  liliick  i>.  '.'ivrii   in   Ki^'.   1(M1, 
8inu8  Arrh}rthmia.     In   this  <-<>iii|itiiiii   the   rixlinl    imlx'   is  imirkcilly 

il  ir'jllliil',  I  lilt   tlic  "m,"  "i'"  ;iinl  "  \  "  W  avcs  nl'  llii'  jlli;iil;ii-  t  r.iiilii;  occur 

With  tlic  iisiiiil  li ril;itiiiiis|iii>  to  one  iiiiotlicr,  iuhI  there  i>,  iio  «li'liiy 

ill  liic  "a"-"c"  iiitci\al. 


:.  uss  ViS 


\  t  «\ix»  A  -\%* 


1  ,c     •.'!.       11,1. umI !:i.IimM    lilii.         I  r  .t    -'  .u.     .i"    I,.  .1,1  l-l-'  ',        'I','        ll      i,,,M.iI-    r,,i-,ll.     !.■ 

Il,.iii    1,  ..    ,,.:,      I  l'i.,'.i    i;,    I'     I  ,111    r,  I 


-S 

I 

I   1 


Sinus  Bradycardia.  The  he.-it  i.ii'jinates  at  hur.'  iiiter\iils  in  ilie 
■-liiiis;  the  "a  c"  iiitei\al  is  normal,  ainl  liie  railial  ]iulsc  \cr\'  slow  Init 
|ii;iclicall\    regular. 

Premature  Beats.  Thesr  mav  he  eiilur  Miitrieiihn-  of  amiciilar  in 
mivin.  Ill  the  I'oiiiicr  e;isc  the  "a"  ua\es  on  tin'  jiiLrular  tiacini.'  .space 
iiLTularlv  thriiii'jiinnt,  Init   tiie  "e"   waxes  at    the    point   ol'  disturliance 


i   1 

3 


'--^WA^-^'UL^U 


'[j.A^y^ 


r^ 


s 


\  s 


r,.-    i,„,     ]> 


I     !,.:■!  !,|".  I,.       ,  I';..-,,     i:.     1'.    e,:'l,  1    I 


I 


'■••iiieiih'  witii  the  "a"  waves,  iiivinir  thei-erore  a  iiioic  pronoinieed  wave. 
Mils  is  due  to  a  preiiiatlire  coiitiaelion  o|'  tiie  \cnlriele  oeeiirriini-  almiit 
t'e  lime  ot'  the  "a"  \\a\(',  so  lli.'it  the  latter  limls  tiie  xentriele  in  a  re- 
"  ,'iiii'.\  sijile  !  sec  paue  i7>!.  'I'iie  pn  iiial  lIlc  coin  raet  ion  is  llieierorc 
i'l'lowcd  liy  a  compensatory  iiaiise,  \\iii(di  is  e\  ideiit  on  the  traciiH^.  An 
■    •iiiiph'  ot'  sncii  a  ease  is  i^ixcii  in  i^'ii:.  Ktl.     In  dniilitl'iil  eases  the  e.xaet 


i 


r^ 

n 

1 

i 

1 

* 

'^ 

h 

.•7M 


Ml     I  IKi  I  I  ATKiN    (If   Till 


lll.tMHi 


sitf  of  oiiiriii  ,.f  thr  |.i..|mitiirr  licnls  i-iiii  l.f  <|r|.Tiiiiti.-i|  ..Illy  li 


Iliritsllli'lllrMi   ..I'  III!'  (|j>t;i| s   Itct 


y  ••ait'tiil 


.1'  th. 


wren  111..  Miiioiis  lioiits  of  t|ii>  vfufrirlt 


ifiir\.T  MM  iiivmilaniv    icpt'iits  it^.h'  nil. I  tin-  diimtioii  of 


nrihvtliiiiia   iicciiratcly   .'ciii.",)) U   to   jtnotluM     i|i 


nno  pvi 


•■    iiiTKiilaiifv 


'"'•         ,'"1  l'"lll,,l„M        :,.,!.       M    ,,,„...,    ,|,,,       w. ,!,„„•..       ,, 

"'     iHl.t\al«   MM.:.. I     1    ,,i,|    /■■•    „,,!,    il,.,.,.   „,,,,,.  .1 


\0  y^\\\  u\  \   '>!> 


t'"    ■'  '■"     t I..ll.-  III.    .illl.lll,..,  .,1 

1  I    /'.      I  l'r..i.i    \      I'    Call.  I   I 


'""•'     '"■   '■'       '"■      ''    I'lviiiiitiiiv    .niri  •iil.n-    or    vcnlH,.,,],,,.    n,mra..|io,is 

'-'   "' ''"-i""''!  urivi, (■  ,|,..,,,|...,|  I,,.,,,,  I,,    ,-„i|,„, v.-iitiinili. 

.•.■>|.o„.s..    :  or  ,:!.   ;,   |,i,.,l,  .|,.i,,,,,.  „,•  I,,,..,,,. 1,1,,,.,,  „i,i,  .,  „|,|,.  variation  i, 

tiic  vrntri-Milar  i-'simmim.      TIi,.  i, rta.it   point   to  not,.  Iuto  is  that    ii. 

tiiHttn-  hou    irn^uMilar  mwI.  a   ,..,,•,.    .    .,„-,.v  a|.,.,ai,  if  ti,..  iri,.;rulu.i1v  ir 
prats  itsril   It  ran   not   I,,'  .jnc  to  auririilar  lil.rillation. 


I' 


,i    1..    '^    t    »,  «. 


Paroxysmal  Tachycardia.     Wl,..,,  tl,..  rat-  „(  a  r.^ilar  puis.,  is  sn  I 


t"  ;■         ;■"•'    ""  " '"'"-'■  '"  '•^""  '"•"■■^  ''  ^"" ''''tio  to  tin.  proviou. 

;■.•"""•";"  ','""'*^"   '"">    '"•  •'"•■  """    l""""itur..  vntririilar  iti.,.- 

t.ons  ^lii.l.  do  not  rcarh  Ih..  radial,  or  r2,   to  ti.o  sudden  dcvrlopnuM.. 


IH>I,VSI'IIV(I>IINIHAMM 


27!> 


,,f  a  twii  uioiu'  hnirtliliH'k.     Wlu-ii  i>n  thr  "tin  r  Imml,  llitn-  is  im  t-xni't 
ratio  li.'t«fcii  the  -»lo\v  iiixl  tli.'  nipiil  nitc.  Ilir  fliaiiiii-  i-  ilili-  In  thi-  n\u[ 


ili>a|>|i<iinniri'    <il"    |ian>\.\  Miial     larli.M-aiilia.      Tin" 

liirh  ilif  aiiriflc  is  licatinn  vfiy  r-a|)i<lly  may  Inst  for 

lilc  linii',  siifli  attat'Us  inisistinL'  "IT  ami  "ii  f"r  li"Uis  "i-  cvfii  days. 


iliii    ap|u'''i'iiii '■ 

pat'oxyNiiiM  tliiriiiu'  » 


.t  \ai'ia 

Till'  tiaciiitf  ill  siifh  a  caw  in  n'wvu  in  I'li.'.  Ill; 


I  IK     In;        A. in. 


(lull,  r         III    till-    '  .!-•■    III!     >•  !•<■ 


I„  'iM    ,„  ,    ,„.i|.,(r 


!■;      P      >    111.  I 


Auricular  Flutter,     It   i^  iiii|i">Ml.l.-  in  .iiiiL'iioM'  lln'  ii"t   infr.iincnt 
.AJslciirf  "I"  this  cardiac  .•"M.liti"ii   \\illi"iil   tlic  u^c  "I'  .itlicr  the  i>"ly 

>|)hy>{liioirraiii  or  tin-  clcdi anliouraiii.     The  .iiliriilar  .urvc   iii.iy   he  "I 

two  tyjvM.  one  inailc  up  "t"  rapiii re  "r  less  iiiiiliMiii  waxes,  the  "thcr 

(if  waves  that  are  pa.ieil  with  a  coiistaril  tiiue  iutervji!  lietwecii  the  pairs. 


t     «-     5 


J^i/^/i^ 


I  1*:     I"4.      Auriuilnr    Hull 


All  of  the  frp(|ueiit  heats  of  the  aurieie  do  ii.  *   •    ach  the  veiitri.-le  in  this 
cnmlitioii.    so    that    the    ratio    lietweeii    a\iri.      ■  ;'    and    \  .'iitricidar    heats 
may  lie  1::!  or   1:4.     The  eonditi.ni  iiiiist   th.'icf.ire  iml   lie  confused  with 
licart-lilock,  the  niaui  point  ol'  distinction  lieine   Ihal    hi  the  i.illci    .  ..ndi 
ti'iii  the  ventricular  pulse  is  slow  and  the  auricular  ahout  noriual.     The 


nil     I  IKTI   I   \ri.l\    ciK 


'I'll!     l:l.(iii|i 


.  i 


l'.'l'li,-|l        |ll||>.>       1||;H         |„.       ,.,.,_;|| 

■"■'■>    ■•'Mii.-iiL,r   l„.,-,i    t,,   ti.nci    1,,   il,,.   \,.|,,,.i,.| 


'I-  iriT!jiil;ir.     Tl 


!■<    |i^ll1ly    Ihr    fr( 


r;h-liii\ 


II'   iMilM'    I'di-   the    r.-iiliii 
'lllliii;,'    ■•|liricMll;i|-    lliil 


t'M\      |i|kis,.     (,r     V  .■||t|i,-lll,-|f     .•nllllJict 


l-l'IKllllnll     ,,|-    III,.     1,1111,11,,,    ,1,1, 1 


Ull'l 


I'll  Illy  111,,  r. 


"'"•      'I'll!'    ImiimIIc    liiMV    he   (•.,iisi,l,.|v,| 


'^■'"^"■"'"    '"■"'^'■"'"'■l'   "illl,:,i,s,„i,   tl„.  i„„H,K,.s  ;„.n.ss  i,   „„lv;.,    ■, 

7'";; ''';'• /'■'''•■'''''••''-•-''•■■-■''--pi.iiv. ,. ...m,..;,,,..:, 

--.1h.  1.11,1..,  ;„„i  ,.S..,I  „f, I,,,.,.  ,|,„  ,,„  .,,„,,,  |„„i„.,,  „„,,,,^^,_,  ^^_^,^ 

n    .■!  ^M,!,,!,!,,,,,    „|'  rx,.i|;,|,ili,v    li.v;iiiso   «(   lli,.    iv 

I''!--    l.-"  .      Tr,i,MiiL:s  slM.ui,,.-  ;iuri,-iil;ir   ilmt.T  ,-,,,■ 


"  ill    till. I    the    \rii|ii,-|,. 
lV.i,-;i.|-\    |,,.|-i 


/-  \  A'\       ,^^      .,1      />^^     r    '      r   >  ,\j-,      e  V         r   V     r  V      t    v    /-   , 


^•e.  wi.Kki.  t)0-.iu,v 


':'■■,  '•''■    \ i^f  -..iMi. 

'■'■■■I    if'L.    1,1. IV     .  ,    ,..,     ,,,|i. 
'""i-       'I-:-  i.     i:^     I'     (  .,,,,  ..I 


1    I-!-..    ..:,  I    l!„ 


"  " ''"     |nyM'.,T-   t,at  nil;,    [I  , 

■'"■I    ••»■■    .l.iMT.j-    l!',     .|.|,M,.,n, 


til''  r.-iili;il   |iiiU,.  is  n../iil;ir 


-'"■"  '"'    '■''--■    I"-!  .-"Hi    ini.      In  „i„.  ,,f  ,|i,.,ii 
111    til,'    ,.l|icr,    ilr,'L;iil,ir, 

Auricular  Fibrillation.     Tl u-.rUuu.  „r  ,l„.  auri..!,..  .s  .hv.-i.lv  .v 

'''•■'""^'l-  •■"■•■  '■"'"•'■l.v  nrr^u\:u:  m,  il,,.,t   tl„.  .jiinul,,.  tnirinns  s|,„u    .-.I'l  m 
'."■'■  '■'  ^^'■';''    "'  •■'"  ■'•■'"  "■•'^"■^'  ""■  '•■•"liiil  ti-,-„-i„._r  Iniii^r  ,.li;ii-,-„-t.Ti/,.,|  l,v 

' "■'".""''''■"■"''^" '■'"I"inii.,i„  rhytl„„.,„|l,y^,,,,,  vMri.lionintl,;. 

'"■'""''"   ""•""'■^"'"■■'1   '"■■"^  I'-i M.  -anli,,.  ,.^,.|,,  ,„,|,„  ,„,„      .,.|,i^ 

:;'"  ;i  -  "<    •!'•■  ^l.M-iiM    i.iis,..     T|..,.,ii..   „■„,„   a  .....  .„■  „„,i,,.L„ 

niiiill.iii.iii  ;iiv  sli,,„n  ill   |.'|._,    |,|- 


<  ii.\iTi;i.'  xxxii 

CIJMCAL  .M'l'M(ATI(i\S  n|'  (  KIMAIN   I'll  VSK  i|,(  MiK  ' 
.MIITIJODS  iCuniMi 

THE  MEASUREMENT  OF  THE  MASS  MOVEMENT  OF  THE  BLOOD 


Method.      'I'lir   ;i|i|)iil;itlls   IIm-iI    t(ir   ll:i>    |Ml|-|insc   c-ii| 
\i'-.scl  ciint.'iiiiini;  ;i   kiidwii  i|iiMi!tily 


;.iiiMI 


i-.i>I>  csx'iilijilly  (if 
w.iIiT  1111(1  a   tlici'- 

iiinirictcr  iVdiii  uliicli  n  cliiiinic  ,,\  iciiiiici:itiii-c  cf  a  liuiMli-cdlh  ni  a  dc- 
i;iit'  cciitiirrailc  can   !"■  lead.      In  di'ilcr  lo  diiniiiish  as  inucli   as  pfissililc 

!ii.'  liiNS  (if  heat    lictwccii   the   \c>scl   and   tl ulsidc   ail',   tlic   waiU   ai'c 

'I'UlMc.  tlic  space  lie!  ween  lieinu'  stuffed  willi  Iiidken  c(ii-k.  Tile  tnp  (if 
'iie  \('sscl  is  cldsed  witli  a  tidck  c(irl<  pkite.  haviiiLT  suilalile  ii]ienin'.'s  in 
It  fill'  tlic  liand  (II-  fddt  and  for  tlie  tlieniidiiietci'  ,-nid  a  stinci-  ifc;itlier) 
'.'.  !tli  wliicli  td  keep  tlic  \\;iler  in  cdnstaiit  nidlidn.  The  apparatus  is  called 
a  iiihiniiK  h  r. 

After  tlie  li.'ind  (if  fndt   lias  liecn   in   the  ealdi-iinetei'.  \\itli  tlic  water  a 
u\\-  decrees  licliiw   that  nf  the  Imdy,  fur  a  certain  time  iteii  niiniitesi,  the 

■■  iiipciMliire  of  the   water   will   df  cdiu-se   |ie( i;iiNed.  aii(l   tln^   deirrce 

I'l  which  this  (.ceurs,  midtiplicd  liy  the  vdliiine  df  the  water  in  ciiliii' 
■  '  iitiineters,  will  yive  in  ealorics  the  aiiKUint  of  heat  dissipated.  Hy  tlic 
.ipplicatimi  (if  a  \-ery  simple  fdrninla  it  is  imw  an  c.-isx  in.itter  Id  caleul.-ite 
ii'iw  iiiiieli  hhidd  niiisi  li;i\,.  passed  thrdimh  the  iihind  xcssels  nf  the  |i;irt 

I'dcr  td  i;i\c  (lilt    the  (iliserved  aiiidiliil   df  heat;   fdr.   if  we  iliridi    thi 

'"/■/i.v    hy    the   (lilhniiri     in    h  in  jii  rdl  •' I'l     hetweeil    the    illtliiw  ill'.'    and    (illt- 
'"wiiiLT  hlddd  (if  the  part,  the  r'csidt   must   indicate  the  rtilimii   nf  lilinnl,  in 

iiiiic  centimeters,  that  has  passed  thi'diinh  it  i  sii 1>\  dclinitidii  a  cahirie 

"i'i''ils  Miluuie  multiplied  liy  dilTereiicc  in  temperature  .  It  reiii.-iiiis  tn 
'  -  I'kiiii  the  e(|uatidii  \>y  w  liich  the  results  are  arrived  at.  If  <>  eipuds  the 
■■i'"i|iit  df  hhidd,  //  the  cahirie-  (if  iieal  ;;i\en  nut  t(i  the  eahirinieter,  T 
-  '  lempcraturc  (if  the  arterial  liludd  and  '/"  the  temperature  ef  the 
•  iiiiiis  hlddd,  then  we  ha\('  the  ci|Uatidii  :     (,)  -     .     It  has  liceii  slinw  n 

1    r   llic    .'i  t<  rmiii  iii-.n    nt    ,'/    we    n<.  i^[    niiiiii[K    (!ir    i:i'iir    i-<  iiti:ii.  t.  i  ^    i.f    \\:i''r    /  '  ■- v    tlir    u.i'rr 

■"■■I    ■■!    111.-    li.m  I   .in. I    ...lu.i.ii.i.r    .l.,,,,ii-,-    i...il,    „t    111,.,     uill    .,l-..il.    -11.,    l„.,n    la    ih.'   ,1,1. 

■■'    t,  ::.;,...:.., .     ,..,.    .:..     -.  a  ,  ...lii.ii:    ...     In.     ,  ,I.n  i.n.i.-i     .i.,,..,:.,-    -mut.     It.  .u    is    |..st    t,,    (li,- 

■■i,,i;    ||„.    ,,|..,iv  ,11.1,11.      Th,     «:u.r    ,,|,:iv.,l,  ,,1    ,.1    III,.    I,i,i„|    IS    ,.,,,,, ,;,    ^,,|,|„„.    iimli  m.Ik  .1 

■:   lli.it   ..I    111.'  ,.i'..,i.r..  t,r  iinisl    I.,-   ,1,1,  ,,■■., I,-. I    e.i    ,  i.li   ir-tril.ll.  til   ,11,. I    I-    ll-,|,illv    .-llftlt    I'tii  ,  c. 
■•   I  .....r,!,;  ,,l    111,    ,.,l,,r,|it,l,  ,    I-   ,i,i,  riiiitt,,!    I,v   ..I,-,  ivif.;   ilf   i.JI    ,,i    1,  i:i;„  , .,1  nr,-    r,,r   ,,   |.,n.„| 
.'    I"    tltlll    ul    lllv    Jlla.ll    ,.:,-ir\,il|MM    Willi, ,111    111,     ll.llitl    111    llii'    ,;i|.,riiii,M. 


I    !|! 


mi 


U 


^l.-.-.!^*^     ^ 


!1      I 


I    \ 


-^-  Tin:  ciKii  i.ATKiN  or  Tin:  iti.don 

ly  Stiuiirt  tliiit  T  iii.iv  Im-  tiik.'n  iis  the  same  as  that  of  the  moiitii,  or  ()..'. 

w  avcrav'c  tcinix'i'aturc  of  tin 


<'.  Ill-low  that  of  the  roi'timi.  ami   '/'  as  tl 
water  in  the  caloriiiictcr  during  tin'  ohsi-rv 

heat  of  lilood,  till    rosult  is  multi|ili>'(I  li.v     - .  tin 
heat  of  Mood. 


itioii.     To  alhi..   for  the  s\h 


reciprocal  of  the  spocific 


Thcor'cticaliy,  then,  the  method 
i\    tech 


is   very  simple,  and   thi'i 


e  are  no  um 
usual  technical  difficulties  in  applying  it.  The  only  si)eeial  precaution 
is  that  the  air  suridumliiiu:  the  ealorimeti'r  should  lie  kept  fairly  con 
st.int  in  ti'Miperalnre.  so  that  we  may  he  enahled  to  allow  in  our  ealcula 
tions  for  the  loss  of  heat  from  the  calorimeter  itself,  this  value  liein- 
olitained  li.\  ohservim,'  th,'  change  of  temperature  in  the  ealoriiiiet<'r  foi 
a  certain   ]ierioil  of  tinn-  at'lei'  the  hand   has  lii'cn   removcil   from   it. 

The  Normal  Flow 

The    i'csuIIn    are    calculated    on    the    hasis    of   ^'rams    of    Mood    llowin- 

""■""-'i  ' ■•'••  "'■  'i^'-ue  in  one  minnt.'.     The  volume  of  the  hand  or  fom 

"^  ■■'■''■'•'•'••ii I  I'y  placing'  it  in  water  contained  in  n  small-si/e.|  irriiration 

can,  the  tnlie  ..f  which  is  connected  -.vitli  a  huretle.     The  heiudit  to  wliieh 
the  water  rises  in  the  Imrelte  is  note,!,  and  after  withdrawins;  the  han.l 
"ater   is  ad<l.',!    from   a    u'raduate   to   the   irrijration   can    until    the   sane 
height    is  reachc.l  (,n   the  Imrelte,     The   iiumher  of  euli.c  <-entiiiicters  re 
'luir.'.j  jiives  the  volnme  of  the  hand.      In  a   imrmal,  healthy  iiidivi.Iu;il 
th.'  averayc  llo«    in  the  hand  is  from    V2  to  1:!  !.'m,   for  the  ri<,'lit  liaml. 
an.l  al.oul  half  a  u-rain  less  (nv  the  left.     This  dilTerenee  hotweeii  the  tw. 
hands  corresponds,  of  eonrse.  with  their  relative  de<.'ree  of  devolopimMit 
The  averairc  toot  How  is  much  less,  and  varies  ac.'ordin;,'  to  whether  th- 
patient  is  sittinir  up  «v  lyin^r  ,|(iwn  while  the  im-asuremeiit  is  heiiij;  ma<lr 
In  a  mniiial   imlividual,  while  lyiiiu'  down,  it   was  n.ll   frni.  in  the  ri-rl.t 
foot  and  r,:2:\  una,  in  the  left,  per  1((0  cc.  of  foot;  hut  only  l>,!Mi  tr,,,.  i',,. 
the  ritjht  ami  4  1  u'ln.  for  the  left  foot,  whih'  sittin-,'  up.     The  results  ha\. 
lie.'ii  t'ound  to  nnderiro  only  a  sliirht  variation  from  month  to  month  in  ,. 
friveii   healthy   individual,  provided  the  aii'  tem]ierature  diiriiiji  the  dii 
feivnt  ohservations  is  the  same  and  the  peisoii  has  l.een  some  time  in  tli. 
room   hetore   the  ol.seivations  are   h.'LMin,      This  precaution   is  especial! 

imi'ortant  if  he  is  a  dispensary  patient  and  has  I n  i„  tl pen  air  wiii 

hare  hands.  The  How  varies  in  ditTei'ent  individuals  Ixith  with  rcfjai. 
to  ahsolnte  amount  and  the  ijitio  hetweeii  the  two  hands  or  feet.  Win: 
the  total  llnu  in  the  Ininds  is  <'ompare.l  with  that  in  the  feet,  a  ratio  >• 
ahont  :;  to   1    is  usually  ohtaiiicd, 

'I  Ic   I'hfisKitofiir  Cnisis  lur  Vnriitl iinis  in  llloixlltnw.      As  ahove  indicated 
the  most    Muirked  of  these   is   prohahly   the   tniijtirnlurr  of  llir   rooiu.      Th 


Pin, 


M 


MCASIRLMKNT    UK    MASS    MOVI  MliNT    iiK    llloiih 


li  inpi  ml  iiri   of  tin   iratir  ni  tin   inliirnin  t)  r  liiis 


lik<'\viM'  ;i  ^rri'iit  iiiriiii'iicc, 


iik)  for  the  i'(iiii|mris()ii  i>(  ditViTfiit  ciisi-s  it  is  always  imimrtaiit  tliat  tlif 
rudiii  and  caldriiui'tcr  triii|ii  ratiircs  lie  stated  aloii<.'sidc  tin'  results.  Miisriddr 
Diitnirtioiis.  iMddiici'd  lix   iiKiviiiLr  iIh'  tiiii-i  rs  in  tin'  caiuriiiirti'r,  cause  a 

(1  tlow   in  tlie  hand 
sir  .liini  iif 


marked   increase   in  tinw .  aeeiinipanii'il   liy   a  diniiriisiii 


that  was  a 
lilt  linn  i>( 


t  rest.    A  trri 


It  <l 


iniiniitioii  III'  lliivv  res'iits  t'rnni  tinis 


sunieieiit  deu'ree  In  olistniet  tlh'  \eniins  eireiilal inn  :  and  w  lien 


the  eon.strietiiin.  as  that  eaiisid  hv 


liliiiiil  |iresMire  armlet,  is  increase 


d  ti 


I'tween  the  systolic  and  diastolic  ]iiess\ni" 


•xlreiiielv  little  lilood  flows 


ihrouirli  the  hand. 

\^y  iiiiinersiiijr  tiic  opimsite  hand  or  tool  in  hot  oi-  cold  water,  the  hlood- 
llow  thnuisih  the  ohscrved  iianil  increases  or  decreases,  respectively. 
The  ciiance  may  he  of  a  teiiiporaiN  chai'aclcr.  or  it  may  iiersist  thronL'h- 
niit  the  wliole  period  of  imiiiersioii  of  the  hand.  'I'hese  rea-tions  are  due 
lo  a  i-<ixriiliir  nlhr.  and  oliservatioiis  of  its  si^nsitix  eness  are  of  value  in 
till"  study  (if  the  elTects  of  lesions  either  of  the  ner\e  or  of  the  ni'rve 
renters  eoneerned   in   \ascidar  rellcxes. 

Clinical  Conditions  which  Affect  the      ^odflow 

l';ven  ill  cases  where  there  is  i)lenty  of  other  evidence  ol  eiirtailineiit 
m|'  How,  the  nn'asnrement  may  he  of  importance  either  for  deteetiiii; 
.111  alteration  in  the  vascular  relle\  or.  liy  "omparisoii  of  the  two 
liands,  for  demotistratinii:  the  relative  dei,'ri  .■  ol  alteration  in  How.  In 
'iriilr  hiUdwwatnrii  niKilHioiis  .■itVcctinji  one  h.ind.  there  is  an  increase 
:ii  How  on  the  afTecteil  side  ;iccompainei|  li\  a  marked  curtailment  on 
the  other  side.  This  indicates  that  an  iiicrc,ised  llow  in  the  infected 
area  is  aeeoinpanied  hy  a  rellex  \  asoci.nstriclion  elsewheic.  i>arliell- 
iarly  in  the  syminetricall.\  placed  part  of  the  iip))osite  side  of  the 
liody.  In  eases  of  nonliaeteri.d  inllanimation  of  the  hand,  as  in  <:ont. 
'lo   siifii   of   vasoeonstrietion    ina.N    lie    oliserved. 

There    are    many    clinical    conditions    in    which    Stewart's    method    re- 

cals  an  alteration  in  hloodthiw    that    wnnld  he  nnsnsiicctcd   hy   the  use 

of  ordinary   clinical   nn'thods.      It    is   for  the   iuvi'sti<ration    of  these   that 

ilir'  method  is  of  <.'reatcst    value.     The   most    imiioitant    lindinyrs   are  as 

fallows: 

Anemia.  The  hlooddow  in  the  hand  may  he  much  less  than  normal 
iM  |iernicious  anemia  and  sccon.ijiry  anemia,  and  distiiu'tly  curtailed 
Ml  chliifosis.  Siiu'c  the  minute  volume  ol'  the  heart  is  al.  o  increased 
•::  these  cnndil  iniis.  the  \  iisocoii-;  li<-1  j.Mi  at  tlie  p'-iiiheiy  \\\\]  r.ssisr 
'u  ciimpellinji  more  hlood  lo  pass  throuuh  the  luiiirs,  so  as  to  make  ui> 
Tnr  (lefioiciiev   of   hlood. 


%^  M 


u 


2X4 


Mr  (  iitii  I  \i'iii\  (IK  -I         iiiiidii 


i: 


Fever.  Siii<v  (■Ii:iii'jc>  in  tin'  (MiIiiiiciius  cii-ciiljiticiii  |iiiiiiiili|y  cnn 
"•''III'''  111"  i-liii'f  iVclcr  ill  III,.  .Ifi-.-iiiiiciiicnt  nl'  the  ti'iM|icr;itiirc-n'i;ii 
hitiii'j-  iiM  c-liiiiiisni  in  \r\,T  i,.|'.  i.:iov  7-'::i.  it  is  ex  idciitly  of  urivjit  ml 
\,-iiil:i'.T   III   lie  iilil,.   til   iiii-n-iiic   >uc-li    cIkiiiiic's   i|ii;in1  iliit i\ cI y.      Tliis   lia- 

' '1  'l"ii''  l'>    >^li'^Mii't   ill  sr\ri;il  c.-iscs  nj'  tyjilmiil   tVvcr  jiiid  ill  line  i-jim 

"''   I'liniliinliiii.      Ill    -niri-.-il    it    \v;is    rnUM.j    tiijil    llii-   llnw    in    the    iVct    Iicmm 

i'\i'i'.i|..|     llic     nnllll.ll     lliiu,    ,•111,1     «;|s    llsll.-llly     II Il     lll'lnw     it.        Tili.S     1(11 

'l'''i<'.^    '"    \''^' n^tlii-timi    V,.,  MIS    til   lie   r:in'i.-,l    iiit.i   i-mu  jilrscriicc.       j-'m 

practiivij     i-ci'.niiv    III,.    Iiainlllow     lias    imt     1 ii    m.    .■Ntriisi\  cly    slii,rr,| 

Tliis  li\  |.Mr\,-ital.ility  (if  tlir  \aMicniistii,.t,,r  mrcliaiiisiii  at  tlir  iicfipli 
I'ly    is    iiiiiM    iialur;ill\     iiit.'i|iivlri|    as    a    di'lViisivc    nai-tini'    nt    tlu^    ni' 

■iaiiisiii    liy    wliii'li    an    inric-isci    siipiily    ,,{    ],] |    is    iiiijinrtril    tn    tlms. 

iiitiTiial   nrv.'ins  «  lii,-|i   l„.ar  tlic  liriuil   ,<(  tlic   iiilVrtiuii.      Wli.'ii    w n 

si. Ill-  that  ill  sjiilc  III'  this  .•niistrictinll  nf  thr  |irri  pliiTy  tlic  lilnuil  j.lTs 
sine  is  liiw  aiiil  tlic  |iiilsi.  liin-ut  ic.  u  c  must  .•(iiiclu,|,.  that  there  is  cni, 
^i'|ci:ilih'    ilil.alatinii    ,,t'   uther    \asciilar    parts,    espeeially    the    splanchnir 

.■MTii.      .\    vrvy   |ii;i.-lic,-il   application   ,.(  these    f;iets   presents   itself   ii n 

si.leiiliL'  the  l-alinnale  n|'  the  enhl-lialli  Ireatllieiit  I'm-  I'ever.  It;  I'nr 
'"^'■'ii'l'l'''    " iiehiije   that    the   eulaneiius   emist  riel inn    is   in    the    iiitei- 

'sts  III'  ;in  ineie;ise  in  the  I  ilnn,  |tl.  iW  til  tile  lll-.^jall  nil  \\lliell  the  stress 
"•'  'li''  iiifeetiiin  tails,  it  will  e\i,|ently  he  iiiiire  ratiniial  to  hiwer  tin 
liiMpciatnre    hy    iiieihn,|s    wliieh    v.ill    net    iliiiiinish.    ami    ni.ay    e\eii    in 

'•'''■•■'■^•''    til''   clltan IS   eiinstiietiiiii    th.-ill    tn   iIm   sii   liy   eailsinir   the   vessels 

'"  'lil.-ilc.      In   other  worils.   the   use  of  aiiti|i\  r.'t  ies  seems  to  lie  eoiitiM 

i'i"'l''''''''l.      il llii'.v    (liliiinish    the    hnily    teliipeiat  lire    liy    ejiiisiiiu'    \aso 

'''''''■''i"'i  •■!'  'I"'  peiiphery  with  a  eniiseipient  w  it  hilraw  al  of  IiIiuhI 
from    tile   seal    of   infeetion. 

Cardiovascular  Diseases.     In  eanli ;ises  the  hamlllou    is  far  mor. 

•'!''    '"    '"■    lliarl-eil!;.     iletieiellt     where    there    is    e\i,len f    serious    illipail' 

iiient  of  the  ni\near.liiiiii  than  in  eases  where  a  umss  val\iilar  h'sim. 
exists  1,1,1  ih,.  hrart  aelioii  is  stioin.;  ainl  or,|eil\.  This  iinlieaPs  tli-ii 
it  is  iMoie  serioiis  for  the  foi-ee  of  til,,  heart  piliiip  to  lie  interfeiv  I 
"itii    than    for    its    \ahes.    pail  i.-iilarly    the    mitral,    to    lie    leaky.      h:\ei 

where    there    is    eolisi.  lei  a  i  lie    \e|iolls    ell  uor.jelllen  t .    the    ||o\v    mav    II,.    lit 

''''  'liiiiinishe,!.  Ill  nntre.iie,!  ,-as,.s  of  iiiirieiil;i  r  lihrillat  ion  the  li|oi„| 
How  is  siiliiiormal.  .\ft,r  th,.  ailniinisl  rat  ion  of  .jieitalis  the  hlooiltlow 
in    siii-h    eas.s    is    oft,  n    |ir,,m|,||\     ami    ,ieeiilei||\     iiiereaseil. 

As    uonlil    he    e\p,.,.|e,|.    ,irh  rinsri,  fnsis    is    assoeiateij    with    a    sm.lll    hln.r.l 

tlow.  ami   the   vasomotor  relieves  aiv  weaker  than    in   normal    persons. 

In  ,,,.,•/;,.  ,,„,,,,■;.„,,  when  !h,-  ;,,„.i|..:.,„  j^  ,,;•  ^,i,.|,  .,  ^•^,.  ^.^  ^^^  ^  ^^^^^^ 
pressiiie  on  the  sulielaviaii  artery  or  vein,  there  is  ;i  ilimiimti,,,,  j,,  |!,,„ 
of  the   eorres|ioii,liiijr   |,;ni,l,   |,i,i    aortie   iiiieiirisin    itself,   altlioii;.'!,    i|    i,i,i\ 


Mi:\srici:Mi  NT  hk   m  \ss   mommint  o|-  iiumih 


,,nis.'  -.'V.-al  .■liiiiiLN's  in  tlir  .-iKiiMMcr  of  tli.'  pulse  ImmI.  du.'s  not  d.vi.l- 
,.,|ly   iiir.vl    tlif   iiiMss   indVcuM'iit    .if   tin'   IiNmmI.      In    : irisin    »(   tlic  siili- 

,-h,viilll    iirt<T.V.    IIm'    IiI.hmU'cW    IMM.V    U'    IMMcIl    -IVilIrr    ill    tiu-    .•ol•lV^|l^ll(\illS.' 

,h„ii  ill  tl;.'  oppn-ilc  hiin.l.  .'vcn  tl  mi-li  the  iiin|.liliiil.'  "f  the  i-iilsi!  is 
\,.rv  (.l.viuiisly  .liMiinishcl  iin.l  tlic-  .lilV.Mvn.'..  In-twcMi  tli.-  systolic  aii.l 
.li.stolic  i.n.ssurcs  dlic  pn-suic  puis..,  is  iini.-li  l.'ss  .>m  lli.'  ntr..'t.Ml 
,han    -m    tli.'   n.iniial    si.l...      I'.y   .mlinary   clini.-iil    iM.-nsiu-..iii.-its,    lli.ir- 

,•,„,.      |-;,is..    .stiinal.s    ,,f    l,l,H„ill,,\v     -AVr    nuWr    lik.ly     tn    I,,'     lllM.l.'.      Til.'se 

,,.Milts  JUT  n..  .l..ul.t   .mill-  l-ailly   t.i   vas.„lilMtMti..n  Unrndit    ah.mf   l.y 

I,,,.ssin f    tl,f    aii.iiris.ii    »u    \\u-    l.ni.-liial    i>1.-ni.s    aii.l    partly    t..    ill.' 

1, ,«,■!■  iTsistaii.'..  1..  til.'  II. .w   ..f  111 1   int..  til..  .lilal.Ml  siil.,-lavian. 

In    l!i„im,u.r.  Jis.ns<.   as    ^^..ul.l    1 Ni....-t...l,    th..    t!..w    is    small,    tin' 

,.„„i,„„i„n    iH.in-    iiM.r.'    .>i-    l.'ss    pr..p..rti.mal    I-    llu'    .lurat  i.m    «(    Hi'' 

,|i.,.,,s,..      Till iitralat,.ral    vasuiiu.t.u.    i...a.-ti..n    t..    .-..1.1    is    als,.    1....-U- 

ii:irl\'   ili1..|iso. 

In  ,i;,tlullr  ,i,n,,in,u   ..f  tl..'  U'ri  tl,...v  ^s  a  v.Ty  sul.ii..MMal  llow  in  l">tli 
■1,.   lian.ls  an.l  tli..  (rrU     Tli."  v,-!-Mni..l..r  r..|l..x..s  aiv  als..  U-Mi-. 

It  is  s..iii,.tiiii..s  .lilli.-nlt  1..  t.'ll  \vli..tli..|.  an  ..hscrvf.!  .■nrtailiii..nt  .>! 
•l,,w  is  a  luTvous  in-II.x  '■f\\-r\  .,r  is  .lu.>  t..  s..im.  MHvlia.ru-al  iiitorf.T- 
.,„.,.  Tli..i-.'  ar.'  t\v..  x\ays  !,y  xvlii.'li  Hi-'  oxact  canso  may  I.o  .lia-n..s...l: 
I  l,v  „l,s,.,viii.4-  111.,  tl.iu  fn.n.  .lay  1..  .lay.  if  it  v,  nains  uiu'lian-.'.!, 
:,iiv  alt..iati..n  must  1...  ,l,.p..n.l..nt  .ni  iii....Iiani..al  ..aiis.'s:  i -J  .  l.y  ..l.s.TV- 
:„.'  tin'  .■liaiM,'..  ill  tl.iw  l.r..nulit  al...iit  l.y  alt..riii..'  tlu'  tcinp(.ratuiv  ..f  tlio 
,,'„u  ..!•  .■al..riiM..t..|.  an.l  s,...iii-  «li..tli..r  tin-  rati..  I...tu...Mi  tli.'  two  Iniii.ls 

., mains  un.-liani:...l   ..r   !..•-• s   alter...].      If   111..   Ia1t,.r   ...•.•ins.   1l,..    m- 

.■inality  in  ll..\v  must  1..'  .In.'  t..  in'rvniis  .•aiiscs. 

Diseases  of  the  Nervous  System.  Tli-.  .IT....!  ..f  onn-lUs  ..n  tlu-  f1..w 
.,.,,.i,.s  Mitli  til.'  .lurati..ii  ...  til.'  .ii>.':is,..  In  .'ases  ..f  .'arly  i..'ripli.'ral 
,„il;,t..val  iH.uiitis  tlu'iv  may  1,.-  an  in.'r.'as.'  „(  \\u^v  alt.'rin..'  ill.'  rati..  I..'- 
■.,r,-u   tlu'   t«..   lian.Is   Willi    th..   ■;r..at..r   ll..\v    ..ii    tli.'   .lisfas,'.!   si.l,'.      In 

■:,.uiitis  ..f  l..!!-  stan.linsi  tli,.  ll..u   is  -•nt  .l..«n.  tli.'  -.M'.'atcr  ll..w urrin..;- 

,1   tlio  licaltliy  si.l.'.     Til.'   .'liaii-rs   lu.rc   air   pn.l.al.ly   .hu'   t..   anal. .mi.' 

.Iirrali.ms  in  tli.'  InnK'H  ..f  tin'  tnl..',  i..'rliaps  a  tlii..k.'niii.r  ..f  tli.'  inlima. 

!i,   nt„l„r-,nin-oii   ilis.os,    \\itli..iit    any    iiiv..l\  ..ni.'iit    ..f  tlu-   s.'us.iry   skin 

:r\rs   th.'    Il..\v   s.'.'ms   t..   r..|iiain    n..rmal    an.l    tli.'    ri.|l..\.'s    t..    I..'   w.'H- 

■laik...].     This  in,li.'al..s  thai    iiiv.ih  I'm.'iit   ..f  th.'  mi.t..r  lu'rvcs  .h.cs  in.t 

.,.,  iVr..  with  1.1 llL.w  t..  aiiythin.i;  like  the  same  <l."-'r.'c  as  iiiv..lv(.nu>nt 

thi.  skill  n..rv(.s. 

Hemiplegia.     A  .!..lii-:.'n..y  ..f  l.l..n,lilo«-  .,f  th.'  paial\/...l  si.l..  is  usually 

■  ..-.I'vcl,  an.l  th.'  vas.miiit.ir  r(.|l.'N..s  arc  altcrc.l.  th..  m..sl   usual  .-liau!;-.' 

II,..:  that  vas..c..iisfricti..ii  is  iii..r.'  .'asily  pi.-.tu.'.'.!  than  vas..<lilatali..ii. 


'      . 


L'xi; 


Till     I  Mill  l.\lln\    UK   Till     lll.onli 


S't 


!?;•    i^ 


m 


III  ■-iiiiii'  (•■■isfs  nil  iiImioI'IiimI   triidciirv    III  \  .Tsnconslricl  iiHi   is  :\  iMiiispiriidu^ 

t'.'alunv 

Tabes  Dorsalis.  'I'lic  lluw  is  ilislinctly  diiiiiiiislird,  pspcrinlly  in  tli' 
ff'ct,  iiltliciimli  uImi  ill  liic  liamls.  ami  ilic  v.-isnniotor  I'dlcxos  ari>  iVolilc 
Sdiiictiiiics  there  is  iiieinijility  in  tlie  lluw  «il'  tlie  twii  liaiids,  wliieli  Ikiw 
ever  need  imt  ne<'essarily  iiidiiiile  ,i  miilaleral  lesion  nl'  the  cnrd  in  tin 
eer\  ie.d    reyimi. 


;! 


•    I      I 


m. 


(•IIAPTKU  XXXIII 


SHOCK 


ShcH'k  n.av  )..■  .lur  1<.  a  variety  of  .•auses.     In  -nuTal  it  may  !>(«  .  .■• 

sri-ilKMl  as  a  ulitiuii  in  Avliirh  tl.civ   is  ino.v  or  less  paralysis  ot   tlie 

smsorv  an.!  .....tor  iH.rtioi.s  of  the  rellex  a.r,  alon-  with  profou...!  dis- 
tml.aiiees  in  the  eirelllatoiy  systei...  s.ll.lior.i.al  tenipeiatiue,  freq.le.it 
Mi<\  shallow  respiiatioii.  ai.<l  iiioir  or  less  u.ieoiiseio.isiiess.  Ceitaui  ol 
iliese  svniptoii.s  luav  l.e  eonsidere,!  as  pri.iiaiy  ali.l  otlie.-s  as  seeoi.-laiy, 
,,„  in.poitant  stei)  in  the  il.vestiiiatio..  of  this  .lifliei.lt  a...l  iinpoitaiit 
l.n.l.leni  beillj,'  to  (listil.U'llish  hetweeii  the  two  v'roUl.s.  Mef.oe  iittel.ll.t 
,.,..'  to  ao  this,  h..ueve.'.  it  will  he  piolital.le  to  .lilTerelit  iate  as  sharply 
^,.  possihle  the  vari.^lis  eoii.liti..i.s  in  wl.ieh  one  or  another  of  the  ...any 
\:irie1i»'s  <»f  shoek  is  saiil  to  ocenr. 

The  following  varieties  of  sl.oek  have  l.e.'ii  ileseril.iMl: 
I    Gravity  Shock.     This  is  eause.l  l.y  the  stai;nati..n  of  hl.io.l  in  the 
>|il:iiiehnie  vessels  an.l  the  eonsecnieiit    ina.h'<inate  tillinsj;  of  the  heait  in 
.lijistole.     It   oeeili's,  when   the  ereet    position   is  assniiieil,   in   animals   in 
vhieh  the  meeha.iisn.  wl.ieh  or(li..ai-ily  eomitensates  for  the  teii.leney  ol 
•,'iavity  to  make  the  l.lood   How   to  the  dejiendent    pai'ts  is  inadequate. 
Ihiis.  when  a  domestieated  lal.hit   with  a   lai-e  i.enduloiis  al.,loi.ien   is 
li.ld  in  the  vei'tieal  lai'  down  position  for  any  leni.'th  of  time,  the  animal 
■gradually  passes  into  a  shoeked  eoiidition  a..d  may  <lie  in  a  shoit  time 
JO  to  ;iO  minutes..     Ohservatioli   of   the  l.lood   vessels  of  the  ear   or   a 
..■,„rd  (.f  arterial  hlood  piessnir  will  show  that  the  eause  ..f  slioek  i.. 
!his  rase  has  been  a  trroat  eurtailmeiit  of  the  l.lood  supply  to  the  up|>er 
,.;nt   of  the   hody.   and   therefoiv  to   the    nerve   centeis    (  Fi'^'.   244).     The 
-boek  is  entirely  dependent  up..u  the  la.\ity  <.f  the  al.dominal  museula- 
Miie,  for  if  a  hinder  is  applied  to  the  ahdomen,  (.r  if  the  exi.erimeiit  is 
iMrfornu'd  on  a  ral.liit  whose  al.dominal  museulature  is  in  siood  eondi- 
!ioii,  siravity  .shoek  does  not  develop.     .Nor  ean   fatal   ii;iavity  shoek  he 
;  .odueed    in    a    dot,'.    altli(.u-h    in    a    deej.ly    anesthetized    animal    a 
iKirked  fall   in   ar.erial    l.hx.d   i.ressure   (.ecui's   when   the   vertieal   tad- 
!..wii  position  is  assumed.     In  man.  in  whom  eonipensation  for  the  ereet 
is  hishly  developed,  shnek  from  -iiraviiy  ..eeiirs  only  wh.en  there 


sture  IS 


l.een  some  other  eonsiderahle  upset   in 
'  also  pajre  24.")^. 


th. 


eii-eulatorv   me<dianisii 


;    M 


i.i 


■    fi . 


,! 


•ss 


■nil    riiic  I  i.\iiii\  (l^■  Till;  iiiiiiin 


ij"# 


•-'.  Hemorrhagic  Shock.     I'n,.  1,.., ■,\,:>.^,.  ,„n,|u,.,.s  ■,   i,v,,i..,|  |j 

ti.>n  or  vlincN.  I, lit   the  ,.Nl..|ii   1,,  u|ii,.l,  ,|iir..|-,    ,1   in.liv   ,|ii;ils  iv;i.'t    l„  ll„ 

''■■'""'  '■''-'' ''  1i«'III"Ii1muc    \;ni,.s  ,-,,„>i,l,.,,,l,|y,      Tllr  .ssrlil  iiil    fiU'U<V   il 

""■    l'l'"IUrtinn    .,!■  llcl.Hinll.'Hii,-   sllnrk    is   .,|'  rullls,'   silllihir   I,.   tllMt    ..f   ITfilV 

il.v  shurk     M.nn.ly,  a  ,|..|iri,nt  ,|i,-istnli,.  (i||i„^,  „,•  ,|„.  |„.;,ri    „ii|,   I,"! | 

"'■'•'"'"   ' '"'"-    ""•   ••^■'■'•l    "f   li.'inorrli.-iu..    \nll    1„.    r,,,,,,,!    .■ls,.ul„.i. 

(  piiLrc  l:;.'it. 

••:.  Anesthetic  Shock.    s„  f,.,,-  .■n  M |  ,,r.ssur,.  ,vil,.x,.s  ,■,.■,.  ...m.vnir.i, 

:'"  ■■.'"""■''  '••■'"  '"■  l^'I'l  ill  ;i  l"NV.-t  r„iHlili,„i  uli,.„  ,.,|„.r  is  ;nliiiinist,T.,| 
111     .illsf     slini.-i,.|||     ;nMn||nt     lu     prn.lll,.,.     Iii.|,t     ;,,„.s|  ||,.sia,  Wlici,     1;,,-,.. 

'Iiiiiiititirs  ,,!•  .thrr  arc  <.|,i|.1m.\  ..l,  a   l>  pic-al  roihlili.m  of  slm.-k   is  aliiins- 

'•'■'■t.iiii   t(,  siiprrvrnr  al'trr  a   tiiric      In   si„-li   iiistan.-cs  llir  art.Tial   lil I 

I'l'sMir,.  n-.iiaiiis  Inw  aiul  -•an  iml  l„.  ,v.tnr..,|  ,.\,.|i  aft.T  an  Imur  or  tu.. 
"T  aftilicial  n'spiration.  Tlicir  is,  liuurv.T.  a  iliinivncc  Ip.^lu.vn  cthri 
Nlinck  aihl    tl,r   \aricly   mIm,-!,    ^^ ,.   sliall    .lis.Miss   later   iin.lcr   tlic   title   m 

Mnri,-al>|„,..L:intlH.  l-nr r,  rr v  al  m'  ,  j,..  anrsthrli,.  .-an-.s  all  r-.tl-vs  I,. 

'vlurn,  uli.r-.as  ni  sur-i.-al  slmrk  ,„„.(  „r  ij,,..,.  ar-  suhM-rnial,     Tl,e  ,|an-M 

"'   •■""■^""•'i-  - I<   liiis  I n   cunsi.i.MaM.v   ,!iiMiiiisli,.,|    in   11,,.  ,-lini.-   \n 

""■  """•'•  '■••"■Hul  a,i,„inislratiu„  „f  hInt  .„■  Uy  tl„.  ns,.  .,!•  ntli.T  ancstliei' 
I'-,  su,-),  as  nitrnns  ovi,!,.  oas.  A  ,.„n,liti„i,  .-Insely  siniulat  in-  sli,,,.: 
'"■'■■  "'""  '"■  ii"lii''"i  ill  til-'  -'arlier  sta-rs  of  tlir  adniinisl  ration  nf  an.  . 
liiHi.s  ul,..n  tli.s,.  ar..  I,a<lly  oivHi,  hut  paralysi.  „(  tlir  heart  ..,•  of  fl„- 
iv>|)iratory  .•.•nt.r  is  a  iisnal  i-ont  ril.iii.iry  i-aiis,.. 

4.  Spinal  Shock.     Spina)  sInm-U  is  pro,|, |   |,y  sreti.^n  of  the  spina! 

'■'"■''■  '""  ''  '^  '"  ' ^•'■Hiilly  .listiniiiiislie.l  from  all  other  forms  of  sl,o.-;, 

Ix'eailse   of  its    |o,.al   eliaraeter.   as    it    alVc.ets   only    tl|..se    p,-,rls   of  II, e    lm,h 
Uliiell   lie  l.elnu    t|i,.  l,.v,.|   ,,f  Il„.   l,.,i„n   in   I  lie  e.nil,      Alpove  this  l(.Ve|   th. 

■■'"'"'•''    '"■•'>    '"■    '"   ■•'    P''M'.elly   no.Mial litioii.   (.xeept    in   eases   Avliei- 

llie   seetion    has   l.eeii   at    so   hi;:li   a    lev,.|    that    it    has   sevi.re.j    the   vasoeo.i 

sliietor   pathway   an, I   tli,-,vl,y    pi„,li |   a    fall    in    hi 1    i.r,.ssnn-    fr,,i,i 

va.,Mlilatalini,.      Kv,n    u  h.'ii    this  has    liapp,.|i,.l    the   iwirt    of  the   aiiinial 

•'""'''■""■  '"  ""■  ■^l''"'il  l'-<i""  i-<  I'.v  I leans  in  a  , lithm  of  sh,.,-!,-.    Thus, 

S|i,.rriiii;t.,ii  ..I...  iv,.,|  ill  a  lii..nk,.y  uli,,s,.  spinal  ,..n-,l  ha, I  he,ii  ent  fa, 
loruar-l  that,  .-ilt  h.,i,!,h   tli,'  p,,st,.rior  part   of  th,.  l,o,|y   uas  in   protoiin.l 

spinal  sh,.,-k   ar,l   th,.  I, ,1   pr,.ssnr,.  v,.ry   l,,u,  the  aniirial   anius,.,!   him 

s,-|f  hy  .■at.-hini;  lli,.s  with  his  han.js.  A  sllfl|,.i,.nt  ,l,.seripti,,n  ,;f  the  e,Mi 
'iition  ,.f  spinal  sh,,ek  has  l,..,.n  .^iv,ii  ,.|s..u  h,.r..,  l.iit  her,,  it  may  I.,.  ii,,l,.l 
that   it  e..nsisls  ,.s„i,i  iaily  in  a   paral.vsis  iiiv,il\  in-.'  at   first   all  of  tli,.  v 

Ilex  m,.,.|iaiiisms,  iii,.|ii,lii|..'  il ntrol  of  lli,.  sphiiK'ters.  in  the  i)art  ,.i 

the  eo,-,!  p.,>f,.,i,,r  ;,,  ii„.  s,-eti,,n.     ill  il„.  ,.,,ui-m.  ,,f  ;,  tVw  ,iays  or  w,.,.ks, 

'"■'■'"''''"-  '"  ''"•  l""^i'i '■  til''  .-iiiim.-il  in  th,.  s,-al,.  ,,f  . level",, pm,.nt.  th.- 

retl,.x,.s  i;ra,|iiall\    r,tnrn.  until   nltimat,.|y  in  a  <.oiii)l '  nmiiths     in  a 


SHOCK 


'JS'.t 


,1,,..    f..r  ..xainpl.-     tliov   i,„.y   M   Iimv.   n.niM"''nv.l.      Tl lU.s.-  ..t    tins 

shm-U    is   11.)   .l..ul.t    ll.c   su.l.lon    ii.t.Tinptioi.    of   tl,r    n.Tv..us    |.Mtluva>s 

«l.irh  ivll.x  iirti .cliiiarily  tak.'s  in  tlir  IiI-Imm-  aniMMi«.  i  s.v  pau-'  SO-H. 

-,   Nervous  Shock;  "Shell  Shock."     «'un>i.l.ral,l..  atln.ti.m  lias  l.-.^n 
,,ai,l  to  tli.>  nrivous  sl.ork  Hull  i,as  fivqurnlly  hr,-u  o1.s,..v.mI  m.  nirn  Ul.(. 

Imv..  i.H.n  sul.i.Tt.MJ  In  tl,.'  lianouin-  siu-lits  an.l  tlu nstant  no.>o  an.l 

„,.rv..  strain   inc'unv.l  in   in.nl.Tn   warlaiv.     Tl.r  sv.nptu.ns   n.ay   appear 

.u.lil.Milv  at  til.'  front  or  tlicy  may  .l.^vlop  in  men  wl,o  liav. mport.'.l 

,l„.,..s..lv,.s  in  an  apparrntiy  normal  mann.'r  until  nMnov-^l  to  thr  roar, 
uhcn  tl.cv  pass  into  a  con.lition  mon-  or  i.-s  si.nulalin-  tiiat  ol  shock. 
Srv.Tc  .-o'lulitions  mav  also  rrsnlt  to  sol,ii.-rs  frn,„  injuries  ^^Uu■U  n.  nor- 
,„al  in.livi.luals  vvouM  .u,t  in  th-.ns,.lv..s  W  Mifti.-i.nt  to  pro,iu.-o  sur- 
-iral   slio.'k.     Tin-   ••liaractcristi.'   sxmptonis   in    -^u.'li   rases   arc   ..|itircl> 

,"lifr,-reut  from  those  of  other  forms  of  si k,  an.l.  as  h.-is  1 n  shown  l.y 

Kliiot-Smith  an.l  T.   li.   I'.'iir.      the  eon.liti.m  must    he  tnat..!   fnuu  the 
murolojiic  .ir  ])sy<'hoi)athie  point  of  vi.'w. 
(;    Surgical  Shock.     It  is  this  vari.'ty  that  is  uoially  vIVrr.Ml  1..  ^vheu 

sp..aks  ..f  sh..ek.     It   may  I...  pr...lue...l  .ith.r  hy  s,.ver.-  m..ehah,.-al 

iuiurv   t..    a    healthy    l.ers.m    ..r    hy    eNt.Misive    manipnlali..n    an.l    r.MO.'h 

hin.liin.'  ..n  the  ..p.^ratiio.'  tahle.     It   is  .■..mm..u   in  tr h   xvartare,  t.e- 

,n„  ,h-n.r..re  an  important   variety  ..f  "sh.-ll   sh..ek.-  whi.-h   t.Mm   u.ust 

l„.  US...1  oulv  in  a  ueneral  s.m.s...      IL.w.ver  |>r.Mlu 1,  th..  sympt.nus  .-I 

Mu-i.'al  sh.M-k  are  very  mu.-h  th.^  sani.'.  The  i.atient  Ins  ,n  a  .pu.^t. 
,,,„l,..tie  c.n.lition.  .•arin..'  little  f.u'  \\hat  is  ..'oin._'  on  aroun.l  Inm.  an.l 
.  M.u.-rin.'  .pi.'sti..ns  only  when  repent.'.lly  an.l  importunat.'ly  .lU.'stn.ii.Ml. 
Hi.  skin.lips  a.nl  -ums  are  vry  pal.'  an-l  un-re  ..r  l.'ss  eyan..tie  ;  the  skm 
;,..ls  eol.l  an.l  is  m..ist  nv  ith  swat:  the  reth'N.'s  are  -rr,..-,tly  .Innnush.'.l. 
1  it  is  usualiv  only  after  applyin-  a  very  painful  stimulus  that  any 
..„„'nt  ..f  .l.'fense  is  elieite.l  ..r  res.^ntm.'nt  is  sh.iwn  ..n  th.'  part  ..f  th.' 
t.     The  p.istural   rellexes  are  als..  aholishe.l.  so  that   if  a   liuil.   is 


,1   it  falls  haek  limi>  an.l  toneless.     The  pulse  at   the   wrist   i 
.1.  thill  an.l  alm.ist  impereeptihle.  ami  the  art.'rial   Moo.l  i 


oressure  is 


lallv  low.     The  resiiirati.ms  are   Ireipiei 


t   an.l  shall. iw.     The  re. 


till  temperature  is 


1     ('.  or  mor. 


li.'low  miriiial.     The  pupils  a 


.lilat.'.l 


iiii'l  I'.'a. 

lii'iit  's  voi.'o  is 


t  slowlv  to  lii-'ht.     When  lie  eaii  h.' 


(liie.'.l  t.>   si).'ak.   the 


honr> 


.1  he  ('.imi)lains  .>f  .-.'1. 


I  an.l  nunil 


mess   111   til.' 


dremities.     The  symi>toms  ar.' 


it  iiiilik.'  those  of  eh.il.'ra. 


Experimental  Investigations  of  Shock 

For  in.luein-  sho.-k   exi-erimeiitally.  two  -eneral   m.'th.i.ls  have   heeu 
eniploye.1:  either  rouirh   manipulation  of  the  alxLuuinal   vis.'era,  ..r  r.'- 


!     i: 


!    f 

iir 


I 


,1i 


If     . 


'!    n 


4 
I 


i'    ■ 


41     i 
It    ^ 


2tM) 


Tin;   ciuri  ivrioN    III'    illi:   lil.dtiii 


lll'iltrll    >tillllllMtio|l    111'    l;il<_'.'    .•llViMCnt     MlTVI'S.       Sjrirc    llic    fNpi'l'illH'Mls    ill. 

llsiljilly   Itcrldi  inc.!   i>ri   iilii'vliicti/i'il   .iiiiiiiiils.   the   clTfct    of   llic  iiiirstlirti. 
liJis  til  111'  iliscMiiiiitcil   in  (•\|n'iiiiiriil;ii  wmk  mi  the  ciiUM's  nf  >liiick. 

'I'lic  liist  sti-p  ill  Mji'li  Mil  invest  iL'.-itiiill  is  niitui;ill\  to  rliissil'y  tin 
syiii|i|iiiiis  inlii  |iiiiii.iiy  iiml  sci-iiijiliii  \ ,  I'm-  mi  tlic  success  iit"  llic  class! 
liciiticiii  iiiiisl  lie,  ■11(1  llic  mitcmiic  (if  liirtlicr  iii\ cstiu'aliiMi  into  tli. 
l>liililclll. 

'I'lic  cai-iici  iii\csti'jatiiis  were  iiMlnrallv  attiadcil  In  fh,  iiniiiiiinin ,i 
full  III  liliiiiil  /ii-i  ssiin  as  llic  |)iitiiary  cause  nl'  slmck.  It  is  tnie  tliaf  .i 
|)iiiiiiiiiiiccil  IdWfijnu  will  ulliiiialely  iniiiliicc  s\iii|(tniMs  tliat  ai-c  nel 
unlike  tliiise  111'  sIh.cU,  hut  on  tlie  otlier  liaml  it  can  icadily  lie  sliown  tlial 
this  is  a  result  milv  a  s\  iii|it.iiii  ami  imt  a  cause  oT  the  emiilitiou.  h 
was  liclieved  li\  fiije  that  the  fall  in  liloud  incssuic  (lepclidcd  on  i' 
uiiivcisal  dilatation  ,,\'  the  Mood  \cssels  caused  Ity  cxliaustion  of  the  ton. 

"''  I'll'  \.i>^ )iistriclor  eciiier.      It  has  1 n  clcai-ly  shown.  Iiowcmt.  th.it 

the  lone  of  this  center  is  practicallx    uoriiial  in  shock,  and  that   the  arte 
rioles  are  niainlained  not   in  a  dilated  Init   in  a  contracted  state,  indical 

in;,'   elearl>    tlicicfoie    that    the    low    hi |    pressure    must    lie   ilcpendeii' 

upon  iiiadc(piate  output  of  hi 1  from  the  heart.     Tlic  evidence  for  tlii- 

eonclusioii  is  as  fojhiws:  (1.  W.  T.  Poller"  and  his  cdjlalKirators  ha\. 
shown    that    hotli    pressor    and    depressor    rellescs    arc    perfectly    unrnial 

in  a  rahhit  that  is  in  a  c lit  ion  of  extreme  slmck.     It  is  particularly  iin 

portant  that  depressor  elTccts  were  still  ohtaiiied,  since  this  indicati  ■ 
that   tonic  activity   of  the  center  must    still    have   hecu   present.      {'2)    Tli. 

hlodd  vessels  ill  a  shocked  animal  are  in  a  trnclcd  state.     (>u  (ipcniiej 

a  vessel  and  iiliscr\  ini:  the  oiitllow   of  hhiod,  an  iiicreas urs  when  tie 

ner\e   to  Ihe  hl.Mid   \esse|    is  cut.      C.',)    This  same    fact    has  1 11  .sliiiwn    h; 

Seeli.,'  and  Joseph."  who  cut  the  vasomotor  nerve  pmcecdint;  to  tie 
vessels  of  one  ear  of  a    while    rahhit    and    thus  caused   a    local    paial.Mi^ 

'I'l'i'ii'i I'  iIk'  vessels.     Intense  shock  was  thi'il  produced  in  the  aiiiiii;! 

Ill  the  usual  wa.\.  after  which  Ihe  hliiod  pressure  in  the  anterior  part  o' 
the  animal  was  suddenly  raised  hy  applying  a  idaiii))  to  the  aliildiiiinai 
aorta  .jiisl  helnu  the  diaphra-jm.  This  increased  hlood  pressure  cause.: 
the  vessels  of  the  deliervated  car  In  hecdiuc  eii!.'iu>;ci|  with  hlood.  hir 
not    those  iif  the   opposite    normal    ear,   which    retained    their   tone    (  I'i.j 

iHtM.       (4.     The    volume    of    hi 1    expelled    liy    the    ventricles    has    he  i 

shown  hy  llcnderson  ■  to  he  distinctly  (liminished  ill  ll jirly  stau'cs  .. 

shock,  the  lack  of  pr iiiiced  fall  in  hlood  pressure  indicatiiitr  t'uil  tlnr. 

must    he    •!    ,.o!i,pens:,!,>!-y    ci.os!  ri.-t  ii,),    i,f    tl„.    ;,  f), -rioles.       Lastly    I."..,    r 

'"'^   ' I'    '"ii'i'l    1'.^    .Morrison   and    llooki'r''  that    the   outllow   df   hi i 

from  the  perfused  oroiins  of  a  shdckcd  animal  is  less  than  that  finm  th- 


v\.i,v.    ,H    iIk     Mi 


'•I    111.     nulil    ...1    1..  IPU    ,lil,,I.>i    1...   .n-     ll,. 
■,    h.~  1.,,  ,,   ,,,1,      .  I-.   ,1,1    V,,  1:^     ,„,|    |,..,|,|, 


,lh,t.,       vU.i.  h 


ill.     .  ..ii»ti  u  r^ 


l| 


)     i 


I 


I.  I 


!  I 


t-'.' 


SHI"  K 


2!»l 


>ilMli'    (iriflllis    llllil"'!'    nnfiiiiil    ri.lMlilicilis.        I 'll  I  I  lliilllMli',     s.vi'iiliu    "f    ill'' 
ii.TNf  111'  siirli  an  iiiu'im  niiiMx  nil   iiii'iiiisiil   millluw. 

Til     tlllM'     NJIlillllS     pilCI'S     111'     l'\il|t  IIIM'     111'     II      inllsllil'll  (I     riinilili-MI      III'     !lt 

l(■.•l^t  <'<'r1niii  i>f  till-  vi'ssfls  ill  >liiM'k.  iiim>    li-  iuMiil   lln'  li'^s  diii'iM   i-\\- 

■  lilicr    t•^ll■|li^lll■ll   l.\    tllC   pilllnr  lit'  till'  stmi-ki'il    lUllirlll    illlil   til"'   iil.lii'lltlnlis 

ihiit    till'    syiii|iiittnlii'    iiriMMiN    mnIi'Iii.    iiistrml    lit'    Ixiiiu'    li;ii!il:ziil.    in 

III    nil   fM'iti'il    stiitc,   IIS   slnivMi    l).\    till'   NNUMtiiiir   iiinl    tlic   dil.iliil    |>ii|iil>. 

I'lirtlii'riiiiiir,    \M'   kiiiiw    from    Itic   ixpciiiiii'iits   id"   Pike.   (Jiillirii'   iiml 

Sti«:irt     '     "HI      till'     IrsllM-itllliilll     III'     till'      lliTM-     I'llllcrs     ill'lrl-      illliTl'iTl'llii' 

with  till'  iMii'iiliiliiiii  til  tlir  Ill-Mill,  tlial    till'   Misiiliiiitiir  iTiiIrr   is  friiiiiik- 
:iM\   rcsisliiiit  t'l  .iiiiiiiiii.     It  i-iiii  w  itlistMiiil  this  .•iiinlitinii  witlimit   liisiiiij: 

its  tiilH'   111-   ITt|r\    iicti\il>     lirtlil'   tlliill    liny    lit'   tl tlifl-  I'Mlililiill    I'rlltiTS 

Tlnisi-  «liii   lia\i'   lii:iilit.iiiiii|    tliiil    il    ililirii'lU'V    in    tlir   toiir   iil'   tin-   vasn- 
I'llllsllirtnr    iiml    iitll.T    lliT\l'    ri'lltl  Is    is    n  s|liiMsili|c     Inr    s| k     ll.-IM'    li:ls,.i| 

tlii'ir  r\  ii!riii-i'  iiaitly  mi  liislniiii-ii-  i-\aiiiiiiatiiiii  ol'  ihtm'  I'i'lls  i>(  slicicki'il 
aiiillinls,  it  li.'iii'.'  assiiiiii'd  tlial  llii'  rlii  .niialniv  sis  s|iii\mi  liy  llnsc  ci'lls 
indiciitis  an  .■NliaiisliMJ  ■■imdil  imi.  'I'iii'  assimiiiliiiii  is,  Iiuvm'mt.  I'litirolv 
iinwiil-ralitcd,  and  im  icj-aid  is  triMii  tu  tlif  \M'1I  I'staKlislird  fact  tliiit 
similar  liisti.l.i'.'ii'  idiaii',;i's  may  in'  |ii  imIihtiI  1i>  oIIht  iM.ndil  imis.  It 
is  (Tfliiiiilv  sail'  111  I'liiii'iiidi'  that  the  rhairji's  in  the  iii-r\i'  i-rlls  in  slim-k 
ail'  till-  ri'siilt  and  imt  thr  i-ai|sr  uf  the  hiw  lilimd  pii'ssilli'  iif  this 
rniidit  inn. 

Since   the    fall    in   aitciia!    |p|<"ii|    picssiii'   hc.miis    with    cmiti  acted    (ir 
Irliulcs.    il    lillist    lie    dc|ii'lli|ellt    "11    n    ilini  iiii^hi  il    il'^i  hnnii    nf    hlimil    friiiil 
11,1    Iniirl.      Iiitclferciice    with    the    heart    aetimi    itself    i  iiii|e|ieiiileiitly    of 
the  lilood  carried  to  this  muan    .  or  a  delieieiie,\    in  the  lilliny:  of  the  \eii- 

liieies  diiriiiu'  i':  stole,     that    is,  a   stasis  of  \<\ I   in   the  \eiioiis  or  cap 

illar,\  areas,  aie  tlie  possihle  causes  for  the  diiiiinished  output.  The 
[■Mssiliility  that  the  heart  aelioii  itself  has  Keen  interfered  «illi,  as  liy 
paralysis  of  the  vavriis  iiieehaiiisi.- ,  ••aiisiiiL'  a  rapid  heatini.'  of  the  heart. 
It.'is  lieeii  shovv  II  to  he  iintenahle     \    various  e\periiiielils.      After  stinmlat- 

iuL'  till iitral   end  of  an   iinciii    vaiiils  nerve  in   the   neck   in   shock,  the 

Il  Ilex  vaL'iis  iiieehaiiisiii  is  still  operative,  l-'nrlhiriiiore,  when  the  arte- 
lial  hlood  ]iressiire  is  artilicially  raised,  either  hy  epinephrine  in.iectioii 
"I  liy  cereliral  ciiiiipressio,i,  the  heart  pidiiipllv  responds  to  the  in 
•leased    hlood    pressure    liy     eoiit  rael  iiiir    more    slovviv     and    vi'.'oroiisly. 

liwdeiitly.  therefore,  ;is  tl ardiae   nil  i-h.-iiiisiii   itself   is  noiiiial,   the   de- 

'i-ielit  discliarL'C  of  Mood  lllllst  he  dependent  Upon  /(///iro/x  r  iliilsliilii 
'  '"1.'/.  .\fter  this  condition  has  set  in.  it  liei'oines  pro-iicssiv  ely  worse 
'■•cause  iif  vveakeiiiiiu  of  the  heart  muscles  i-onsenilelit  upon  the  failini; 
'   '"ill  siifply  tliriiUL'h  the  coronary  vessels. 

I  hi    (fill  •iliiiv    lliifi  fun     iiiirriilfs    ilsilf    limni    In    tin    innsi    iif    llii     iiujji- 


1 


ll 

is    Is    I 


I    t 


292 


I'lir  (  ii{(  I  I  \riii\  (IK    iMi:  iii.doD 


h' 


'  i  \ 


i'-i 


'1 


runt   rdlir.     ul    n  inms    hhniil   lo   tin    hiiiii.      Ill    tlio    lirsl    liliicP,    let    US  sor 

ulictliiM-  sliiM-k  rjin  III'  iirniliirc'd  fx|iciiiii('iilnllv  in  niiiiii;ils  liy  nifclianicil 

intcri'croK-i-  witli  tlio  iil lilnw   in  tlii>  xcna  f;i\;i.     That  such  is  the  Oiisr 

was  >li(i\vii  liy  II.  II.  .iiiii.'wav  ami  .lackMiii.  '  \\\\„  t'oiiiid  that  nicclianical  (ili 
struct iiiii  III'  till-  iiiliTiiir  vi'iia  rava  {<>v  a  simrt  tiiiif  was  rnliowcil  liy  tlio  usual 

siiriiN  III'  >liiii-l.;.     Siirh  iiitcii'i'iT witli  tlif  \fiiiiiis  ft'tuiii  t(t  the  lioarl 

iiia.v  alsii  111'  |iiiiiluci'(l  liy  lArosixc  iiiii\ niiriils  III'  tlic  tlicira.\  as  a  iv 
suit  111' artilii-iaj  ii's|iii'atiiin,  Tliat  this  in  itscil'  iiia.\  i-ausi'  slmcU  is  Uikiwii 
to  all  f.\|>(  riiiinital  iii\  fst  ii^atm  s  mi  ihr  siilijiM-l,  allhuiiLrh  the  iiitcriirc 
fatiiiu  has  imt  al\\a\>  Ijith  that  which  is  i.'ixcii  ali(i\c.  Yaiiilcll  Men 
iltTsiiii  -  tlidtluht  that  the  c\ccNsi\c  \  cut  ilat  inn  cailscil  a  lilii«ili<;  ntV 
111'  cai'lii.ii  i|i(i\ii|c  I'l-i'iii  the  lihiiiil  I  sec  |iaL:c  '2'X'i'.  thus  iiriuliiciiii;  a 
liiw  tciisiiiM  III'  this  "as  ill  the  liliiml  (  acapnia',  \\liicli  he  liclicvi'il  ti>  he 
till'  rcspniisililc  lactiir. 

As  ill   i:ia\ity  si k.  mi  in  siiruical   >hiick.  .sIik/ikiHuh  nf  hlooil  in   tin 

.^lilitiirli  iiic  mill  is  riiiiniiiiii :  the  animal   hi Is   into  his  uwn    i  s|ilaiichiiic 

IiIiiikI  vi'sM'ls    1  ca|iillalii'-N  ami  M'linlcs   .  Iiccansc  iIicm'   have   lost    their  tune 
.\s    We    ha\e    Hilled    aliii\e,    iiiie    iil'    the    iiiiiNt     eiTtaill     w  a.\  s    ill'    |iriiillieillij 
-.hiiek  i>  hy  e\|inNuie  and   rmiiih   handlinir  nl"  the  aliihiiiiinal    viscera.      It 
is    tlieier.ii  e    n\     iiilliiiltance    tii    study    the    etTeets    that    can    lie    lliited    mi 
the    liliMiil    Vessels  iil'   this   a  lea    under   slleli    eiilid  it  inns.       Wlieil    the    viscera 
are  liist  e\|iiised  In  air,  theie  ma>    he  a  slmit    |ieriiid  diiriii!,'  which  vasn 
eniislrictinii    is    e\ii|i'ii1.      This    is    sunn    rnllnwcd    liy    a    dilatatinn    nt'   the 
arterinles  in   the  e\|inscd   ana,  ciiisinu;  the  caiiillaries  and    veins  tn  he 
cnme  markedly  distemlcd  asdiirine-  the  lirst  staiic  nl'  intlamniatinn.     This 
aeeuiliulatinii    111'  hlnnd    ill    the   liiesentelie    veins  lias   lieeii    shiiwil   liy    .Mm 
risnii  and   Hunker  In  cause  all  increase  in  the  wei^iht   nl'  an   isnlated  hiii|i 
nl'  intestine  as  an   .'iiiiinal    passes    inln  ;i    stale  nl'  shuck. 

Splanchnic  eimm  uenient  ,'iliine  i|nrs  lint,  liiiw  c\  er,  siii1ii-i'  In  cNplaiii  ali 
the  hiss  111'  liinmj,  and  we  are  driven  In  cniielude  that  the  capillaries  ni 
the  tissues  mitside  the  ;ilidmi|en   liillst   cntr.ip  much   ul  it.      As  a   matter  nl 

I'act.   ('aniinii.   ami   nllieis,    |,a\e    I'mmd    tli.il    enneent  lalinn    nt'    the    hi I 

-urs  in  these  eapillaiies  as  indicated  hy  cmiip.irisnns  ul  the  enrpuscles 

and  heiiiiejlnliin   in  lihiml   diaun    I'rnm    veins  and    I'lnm  capillaries.      .\nr 

Iliall.V    the     V.llues    are    eipial  :     In     slmck     the    c;|pi||;il\      lilnml     is     much     cull 

ceiitriited. 

In  so  I'ar  as  the  eireulatnry  disturhanees  are  enncerncd.  we  tnav  there 
flire  Slim  up  the  cmiditinns  nccurrinir  in  shnck  as  rnllnws:  The  lilnn.! 
aceuniiilates  in  tlie  veins  anil  capillaries  that  is,  in  a  |)aft  <if  the  va- 
c'uliir  system    that    is  lieynnd    v  .•ismmitnr  cnntrnl.      The  cnnsei|Ui'iit    with 

drnwal  nl'  this  hi |   I'rnin  the  cireulatinii   prndiices  a  diiiiimitidn  nt'  lie' 

lilnodllnw    ill  the  Vena  cava  and  cnnsripiently  an  iiiailei|uat('  (illiiiL'  nt"  lie 


\%' 


SHOCK 


293 


hciii't.  The  coiiseqiit'iit  fiutailini'iit  in  flic  >vstiilic  (lischiiryc  docs  not. 
huwover,  at  first  caiiso  jiiiy  ninikiil  t'iill  nl'  arterial  iilood  in'cssurc  1)0- 
i:msi>  of  a  i-cci|)i(ical  coiisliict  idii  cif  tin'  iici'iplicral  aitcrioles  of  llii- 
liody.  .Mcaiiwliilc.  lioucvcr.  the  sta^riiatioii  (d'  iilood  in  tin'  rapillary  areas  is 
iniiiricssively  irici'easiiiji:,  so  lliat  less  aiid  less  Iilood  remains  availalile 
I'.ir  the  syslenne  eireulation.  ('onse(|Uently,  al'tei'  a  while,  in  sjiite  ol' 
ilie  arterial  e msti-ielion.  llu'  hlooil  pressure  falls  to  the  d,in<j:<'roiis  slioek 
le\cl,  and  the  secondary  synipt'H'is  of  fall  in  temperature,  (lulling;  of  the 
rellexes,  etc.,  supervene.  Inei-easini,'  \isc-idity  of  the  Mnud  ;dso  retards 
its  flow. 

The  fundamental  ipieslion  in  the  pathogenesis  of  shock  cimeeriis  tliei'c^ 
fi.re  ///(  cinisr  tif  tin  stiiiiiiiilinii  af  rii-cidnldrii  lUt'nl  in  /A,  (  njillhirii  s  <uiil 
ri  mills.  Two  hypotheses  havi'  lieen  .ilfen'd.  oni'  heiuL'  that  the  stiliudatiou 
III'  alTi'rcnt  iicr\  c  fihei's  to  the  i-cspiratory  centei'  causes  c\ee»i\  c  ah  eolar 
\cntilatioii  with  a  consei|Uent  \\ashin;;  out  of  carlioii  dioxide  from  tin' 
Mood  lacapinai,  which  cnuses  a  \  eno-capillar>-  atonia.  and  the  other, 
lli.it  a  hoinhai'ilment  of  the  vasoconstrictor  and  oilier  nerve  centers 
li.\  alTercut  impulses  liiiiiiis  thoe  center^  into  a  condition  of  exliaus- 
lioii.  which  is  the  essential  cmusc  of  s||,icl<.  The  .-icapnia  h.\  poth- 
'sis  iiniy  lie  ;it  once  dismissed,  since,  on  the  one  hand.  il  has  liecn 
shown  that  in  t.vpical  ^\un-V  there  is  no  delicicncy  of  cu'lion  dioxide  in 
till'  venous  Iilood  (Short'.  ■  ;ind  on  the  other  hand,  conditions  of  shock 
.■i!r  often  jiroduced  without  excessive  In-catliini:. 

\oi-  is  Ihi'rc  an.v  ev  icicnce  to  support   the  v  iev\    that  si k  is  caiiscil  liy 

t':itiu'ue  of  till'  cardinal  centers  us  ;i  result  of  excessive  sensory  stiniti 
l.-iiiiii.  Ill  the  first  jilacc.  it  has  liccn  shown  hv  Mann  '  that  diiiaiiv'  han 
'IliiiL'  of  the  ahilominal  viscera   thi'  nervous  impulses  1  raiismittcd   up  the 

■spinal  cord  arc  much  h'ss  marked  than  those  traiismilted   when  th n- 

n,tl  emls  of  sensory  nerves  .-ire  stimulated  l>.v  opcr.itive  processes  on  the 
Hiniis  ;md  joints,  jillhoimh  shock  is  much  more  re.idily  inoduced  \i\  the 
lnriiier  procedure.  The  melhod  emphi.ved  liv  Maim  for  delect  ine-  the 
i'\isteiice  of  these  afferent  impulses  w.is  ihal  of  rmlies  and  Miller,  in 
''liich  electrodes  are  plai'cd  on  the  lir.iin  stem  in  deeerelirate  animals, 
:!!iil  the  current  of  action  which  accomp.iiiics  ihc  pass;iij:e  of  nerve  iin 
[I'llses  rctristered  h.v  a  strinir  i^.il v  ;iiiomet(  r  .Mlhoiiuh  this  method  is 
^■inplc  and  ilircct  in  principle,  il  h.-is  lieen  found  hv  .Mann  to  ie(|uirc 
-Ti  at  cjire  in  practice  liccaiisi'  of  the  fact  that  the  sjiehiest  movement 
'  the  head  end  of  the  animal  prodiiies  dellcclioiis  u^  the  e.-ilv  anomeler. 
!■  the  further  results  of  this  iiiv  cstient  ion  should  show,  as  the  earlv 
"ii"s  have  done,  that  shock  iiia.v  lie  produced  in  an  animal  withoni  an.v 
■liserved  dctlcction  of  the  u'alv  anonieter,  it  will  disprove  once  aii<l  for  all 


11^ 


il'l'S 


M 

m 


i 


•JIM 


Tin:  (  ii{(i  lATiuN-  OK  Tin;  iii.oun 


till'  tliciiiv  llint  sliiM-U  is  ili'|ii'iiili'iit  M|ioii  iiri  iiii|i;iii'iii('Mt  ol'  a  hiirlifi 
iiiTM'  iiiiM-liiiiiiMri  as  M    ii'siill    (if  civciNtiiiiiilalicpii   l.y   alTcicnt    iiiipiilscs. 

t'aniiciii      lias  i< ntl\    su-'icslcl  that   tlii'  cii.ucni.'i'iiiciit  of  tlic  sjilaiicli 

iiic  IiIcxmI   \(-sc1s  may   he  the  irsiilt    (if  a  fniislricli.iii   of  tlic   |)..i-lal   I'a.l 

I'-lcs  ill   the  lixcr,   uliirli   daiiis  liai-k    tin'  lii I    in   th,.   |iiii'tal   cii-ctilation 

lie  |iiiiiils  iiiil  that  llicsi'  railiclcs  lia\('  \  asdcuiist  liclur  iii'i-\  c  (iiicis.  a^ 
.■\  ill, ■nerd  liy  til.'  fact  tliat  llir  late  cif  ||,iw  iif  Iliii<|  tlinmi;!,  |||,.  |„.r 
liisi'd  liver  iln-rrascs  iliiiiiiLT  aspliyxia.  as  well  as  wlicii  the  lii'patic- 
iirivf  |ilc\Ms  is  sliniiilalcil  cliM'trii-ally  m-  w  licii  cpiiirpliriiic  is  in.ji'ctril 
iiitd   the    piiital    \ciii.      He   aiLMics    llial,   since    tlic    lilnn.l    \csscls    in   ntliir 

aicas  cif  the  IiimIv    all'  enlist rictcil   in  si k,  s<i  also  will   lie  lliose  of  tlic 

liver.  v\itli  the  result  that  the  liloml  of  the  pmlal  vein,  in  uliieh  onlinarilv 

there    is   a    very    low     lilooil    pressure    I  III    iiilii.liyi,    will    liecoine   d: ed 

hack  in  these  veins  and  iherefore  reiiioved  from  the  svstemic  cirelllatioii 
it  does  not  seem  to  the  writer,  liowever.  that  this  cNplanation  is  likilv 
'"    '"     'Ik'   eonect    one,    for,   althiin'_di    it    is   true    that    vasocoiist  rietor    in 

lliiciiccs  have  1 n  shown  to  exist  in  the  hepatic  radicles  of  the  portal  vein. 

.^'■'-  "^ii i'    i^  "Illy   lindei'  special   expelimeiital   coiiditiniis   that    this   cm 

I"'   'I 'hi'.v    liiilst    lie    wry     fclilc    in    nature.      As    we    have    seen    else 

v\  here,   portal    v.isocoiistrictiiin   can    not    he  deiiioiist  rated    liv    stimulation 
of  the  hepatic  plexus  with  stimuli  which  are  sutlicient  to  iirodilcc  iiiarked 
coiistricliiiii  III'  the  hepiilic  ,'irteiv    radicles  i  see  |iac-e  I'.'i.'i ) . 
Till'  eni:oi-i:emeut    of  the  splanchnic   cjipilljnies  and    venules   is   much 

'"'    likelv     to    he   dependent    Upon    liiiiil    i  ii  ll  iii  iiri  s   nclliii/    ml    tin     riss,!-. 

Ihi  HIS,!,;  s.     When  shock   is  produced  liy  manipulati f  the  alidominal 

visceiji.    it    is   easy    to    see    how     this    local    distiirliaiice    mi-jlil    lie    set    up 

\^l'''ii   ^1 k   is  caused    in   other  wa.vs,  as  liy    violent    stimnlati f  sen 

sor_\  nerves,  the  jitonv  of  the  spl.-mchnii'  vessels  is  not  so  easilv  accoiiiileil 
for  unless  \\v  assume  that  il  is  a  t\  pe  of  .ilinoriiial  reciprocal  vasciila' 
innervation,  j'or  example.  v\  hen  stimuli  ai'i'  applied  loc.illv  to  seiisorv 
snrl'aces  under  ordinary  eniidiiioiis.  a  ilist  riliiition  of  the  lilood  of  tie- 
hoily  takes  pl.ice,  mote  heiiiu-  sent  to  the  irritated  rej;'oii  and  less  to 
Ciller    parts    of    ihc    hodv     •  see    p,|._re    -.'.Isi.      Duriiiu-    the    sensory    stimiila 

lion    piece, IJiii,!-   s| k.    il    is    , ivalile    that    this    reeiproc,-il    iiinervat ion 

acts  in  a    faulty   nninner.  caiisiiiir  at    first   a  dilatation   of  the  splaiielinic 
arterioles  and    thus  allow  inu   more   lilood   to  enter   the   splanchnic   capil 
laries   and    venules,    which    lieinL'    possessed    of    litth'    tone    are    inca|ialil' 
of   respoiidinji'  hy  increased   tonicity,   so   that    they    liecome  overdistelidcl 
and   the   lilood   ill    them   stau'ii.'itcs. 

In  anv    ease  there  is  no  douhi  that   the  initial  clianue  is  the  stai:iiatioh 

•'•'   I'l I    ill    these   vessels.   :ind    when    once   such   sla'iliatioli    has   nceui  red 

(lie  inocess  <t<\vs  oii  spniitaiicniisly  ]nolialily  on  account  of  (he  acciiiiiiila 


I     ': 


1 


SHOCK 


i!>.1 


linii  ill  llic  slii(iii;iiit  lilciixl  III'  iiii'oiiipli'ti'ly  tixidi/cil  iiictiilicilic  prndiicts, 
\'.  liirli  liiisc  the  li.\  (linircii-iiiii  cipiirciiliiitiim  (if  tlic  IiIiiimI,  iiinl  priKliicc 
a  I'lirtlu'r  ri'l;iXiitii>M  nl'  tin-  Miiisclt'  lilicrs  in  tlic  vessel  xv.ills.  Tli;it  iiciil 
his  siicli  .III  elTiM-t  is  well  known  (  pii'jc  !t:!T).  Diliiliition  or  .itonieity  thus 
luimressiv  ely  iiicrenses  iiiiil  is  nieiiiiwliile  I'liillier  eneoiir.iiieil  liy  llie  <|e- 
priN.ition  of  o\.\  iren,  for  it  Inis  lieeii  sliow  n  tlijit  isolntecl  jiftery  strips  do 
impI   exliiliil   liicir  nsiiiil   toiiieitv   when   deprived   of  o\y;;eii. 


Treatment 

Wli.iteMT     |n:iy      lie     tile     cMUSi-     of     tile     ;ito|iy     of     llie     eMpillaries     Illld 

■..■iilliev    ill    siioej^.    the    (Xisteiii-e    of    this    i lilioii    ilidic'ltes    tllMt     tl st 

|i..|icfnl  line  of  t  i-i'iitiiienl  is  to  eiinse  the  vessels  to  re;iei|uire  their  tinii-. 
It  uill  he  reiiieiiilicrecl  th.-it  in  irnnilv  slioel<  in  ;i  r.-ilihil  recovery  may 
Im'  aeeiiiiiplishe.l  liy  the  applieal  ion  of  a  ti'.'ht  himler  to  the  aiidonien, 
.ii  hy  plaeiiii;  the  animal  in  a  head-ilow  ii  position.  Siieh  measures- ap- 
plied in  the  ease  of  man  have  ,io1.  liovvi'vcr.  Iieeii  foiiiiil  of  miieh  value. 
I'icsMire  thus  applieil  is  evidently  not  liroin^lit  to  hear  siiflii-ieiitly  on 
Hi''  atonie  vessels.  Cannon  has  theret'ore  made  the  iiiterestiinr  suL'ii'estion 
th.il   a  hopeful   iiioecdiire  nia.v    consist   in   inJectinLT  directly   inio  the  al)- 

■  i. linen   a   saline  solnti •ontaininy-   piliiitrin,  a    hornnnie   vvhi(di.   it    ^vill 

!ii'  n'liiemliered,  acts  direetiv    on  involuntary  muscle  liher. 

Tuo  other  methods  have  heen  ailvoeated   foi-  the  treatment   »\'  slnnd; 
iiiiiely,   saline   nf   Mood    transfusion    and    injection    of   epinephrine:    but 

•  ilher  method    lias   proved   of  practical    value.      Kpineidirine    iii.jei'lions 
i!n  indeed  temporarily    raise  the  arterial    lilood   jiressure,  hut    the  suhse 
1  i|int   condition  of  shock   is  iiossilily  vMirse  than  that   oiisiinally   present. 

\rtcr   the   injection    of   Mood   or   saline   solution    containiiiir    i;elatin    or 

■I ilau'e.  the  hlood   pressure,  althouiih  temporarily  raised,  vcrv    ((uiekly 

i.ills  au'ain.     In  this  re<j;ard  snru'ieal  si k  dilTers  from  the  si k   follovv- 

iij  scvci-e  hemorihaire,  in  vvliieh.  as  explained  elsevviiere,  recovery  cd'  tiie 

■  ' 1    pii'ssui-e   as   vvi'll    as   of  the    ireiieral    coiiilition    of   the    animal    can 

•i    ; iiiiiplislied   liy   transfusion   with   Mood  or  with   saline  solution   con 

liiiiii'-.'  mueilajre  or  "iclatiii.  Tliis  would  indieati'  that  there  is  some 
■^^iiilial  dificrence  lietwi'cn  these  twn  I'oriiis  of  ^liock  i  see  \i,\'/f  14(1). 
llie  only  tri'atmeiit  of  avail  appears  to  lie  to  keep  llie  patient   warm  and 

■  1  leiinive  causes  of  excessive  pain. 

Causes  of  Secondary  Symptoms 

I'  remains  to  einisider  the  cause  of  some  of  the  second.ir.v  conditions 
I  lopiiuj  ill  slioek  namely,  the  ilisi iirlniiiri  s  in  si  iisnlioii  mnl  iiKitinii 
■   llti   full  ill  boihj  Ii  win  nil  lire.     All  of  tiu'se  are  uiiilouhtedly  deiieiid- 


I     , 


;■    I: 


'it'  ^ 


29<i 


■nii;  nui  t  i.ATio.v  of  tiii;  lu.iMtK 


flit   uimu  tl„.  |„u    i.itcii:,!   1,1.„.,1   piessiitT.  Mltlu.Ufili   somil"  authors  l.av,. 
su-'cMcl  thai  II, ,.  !,,»  ,,iM.nsali(.ii  mav  \,v  .h.|K.|„hnl  iipnu  an  iii.-iTas...| 

_'''■'".""'■" ■    I'I'M-k    ;it    til,.    >.M,;,|,>,.s   ,,r    th.'    ivrrpt,,!-    iiciirous    ,  pa^'C    S(i:;,. 

Tliis  Mi-trotinn  <h|.fn.|s  on  the  fact,  .linmiistialfd  In  Sliciriii!,'t..ii.  thai 
iv|>-.itr.l  vliiiml.iti.,.1  u(  the  ic,.,-pt.„s  of  a  vv\U-<  an-  pr.Mliuvs  fafiunir 
"I  tli.il  parti, ■iilai-  ivilrx,  an. I  that  this  fatifiu..  nnist  1...  ivM.h'iit  in  1h. 
N.Miap.sis  ami   not    in    the   niut.n-   n.-mun.   sin.-r    ih,.   Nan..-   iih.tor   n.niiui, 

that  paitiripat.,|  i,,  ,|„.  f,-,ti;;n.'  ,-aM  still  1 I|,',|  into  activity  ],v  alVcrct 

MM. mil  tianMMittcl  to  its  ncrxc  c,.||  thn.n-h  other  sensory  pathua.\s 
IS,.,,  p,,^.,.  SL>,-,  I,  i,  t||,,||„i,,  ti,;,,  j,|  ^|„„.|.  ,1,,,  iv,,,|,|,.„t  jiiVcrcnt  stiMiiiLi 
tion  pro,|nc,.>  synaptic  fati-uc  ahd  tiicrcforc  , lulls  the  seiisorv  responses 
of  the  animal.  The  ,  esiarejo's  of  M^nn  ahovc  referre.l  to.  i'n  wliicl,  \u- 
sIm.us  that  sh.M^U  may  oc-nr  with.Mit  any  .Icnioiistral.le  afiVrciit  .stimuli 
ml  lie  hrain  .t.  iii.  v,,,nl,|  seem,  liouever,  to  neu'ative  the  ahovc  hypc.thesis 
Tile  iai-e,l  thi-eshoh!  of  sensory  stiiiiiilatio;i  is  no  donlit  an  elTcct  of  ih,. 

'""    ''' '   prisMire.   It  lias  l„.en  sh,i\vn.   for  .'xamph',   l,y   K.   I,.   IN.rter - 

tliat  «l„.n  th,'  art-oial  h!oo,l  pivssiirc  is  mainlaim'.l  at  a  uniform  lev.'!, 
til"  liir,si,„l,l  stimulus  f,,r  spinal   ,-or,|   ,vlh-\,'s  remains  i.ractii^all v   uni 

''"'"'•  '""   ' """^  I'romptly  iiHMvas,.,!  xvl„.„   ,|ie  ;irt,.rial   liloo.j   pressure 

1-  ina.h'  t,,  fall.     \Vh,\   a  h.xwr  1,1 1  pressinv  shoiihl  hav,.  this  elT.'et   is. 

lio«,.v,.r.  ,|ifli,.|i!t  to  uii,|..rstan,l  in  tlm  li,.'ht  ,,f  the  r.-s,.ar,-li,.s  ,,f  St, ■wart 
.111,1  his  ,-„u,,rk,.rs.  ^^\uK  as  r,.mark,.,l  ahovc.  foun.l  that  the  cells  of  tli,. 
ivntral  n..r\,,iis  syst,.m  may  emlnre  t„tal  an,'mia  for  maiiv  minutes  an, I 
still  r,.,-,,v,.r  tl„.ir  physi,,|,,^M,.  con,liti,,n.  It  may  he.  h,,uVver,  that  tie- 
l,,\v   M,io,|    pr,sMi,v   .-iffi.i.ts   Ih,.   c,,n,liicti\ity    ,.f   tli..   synapsis. 

Th,'    miiM-iilar    xwakncss    is    prohahly    als,,    .h-p,.,,,],.,,!    „„    |„\v    1,|,,  i,| 

IMvssniv.   f,,r  it   has  l„.en   f,iiin,|  in  animals  that,  wh.'ii  the  art, .rial   hi I 

pressure  is  |,,u..,...,l  f,  ah,, ill  !M)  mm.  I|,j.  tli,.  mus,.h.s  .M.ntract  miicii  l,s. 
•■fli'-i.'Mtly  than  or,linarily.  The  fall  in  hoily  temiM.ratnre  is  ,l,pcn,l,.n. 
upon    thi.   miiM-iilar   iii,.|'li,.i,.n,-y. 

In  e,,ii,.lusi„n.  it  sh,,ul,|  l„.  p,,i,„,..l  „ut  that   W.  T.  I',,rt.r.  in  tli,.  im,- 
"-■"'""  '"■  '"•'"'■  -'""-'^   ""■'    "itii  at   th,.   fr.uit,  has   f,,iin.l   that,   in   manv 

i-as..s  at    hast.  i|„.   ,-i,vul,-it,,ry   .jistiirh; ,■    is  ,ln,.   i,,  a    c-m,|iti,,n    ,if   fat 

'''"''"''"'"•  '''li''  i'^''  i^  ileriv,,|  fr,,m  ti,,.  niarr,nv  .if  l,,im-  |.,,n..s.  siu'li  a.- 
th.'  feiiMir.  I,y  i,,J,iri,.s  «hi,.||  Miia^h  th..  I„,n,.s.  l',,rt,.,'s  ,,hs,".rvati.,n. 
arc  at   h'ast    \  ,.rv  su._'i;',.sti\  e. 


<ll{<  IT,.\T!<t\  HKKKI.'KNCKS 

I  .M,iii,i^'raplis  ) 


\\  i         ."  <      (  '        I     ■  'jM,,      *  ■; .     ..I.  . :                     1  >        ,,  .               ,      I .  ■ 

,,  -,-  ■  ,•    ■   ,  '    '    '  '■'    "'"■•'"   ■■■■'■'■'    l'-.:.M..    I-I,, I,, .i,  ;,,;,,:,.    I;,!,-,. 

M:Hk., ./,.■.  .1.:  1.,,,:,.,.,  ,,,•  ,|„.   II,.,,,,    ,,M,,i-,|    .\I,.,|i,..-,|   |'„l,|Ul„.,.^    ..,1.   -     ,.||,, 

'•'■"''■"';'"■■'■■  -^l-l'^'i'^"    "I     Hie     II. art     l!..:,i,     lull.    sli:,u    .>e    Sen.     K,.tt,r    I...,.,... 


m 


SUOt'K 


297 


Hi 


t  : 


l.iwis,  TliuiniiM:      Iliirvi'v  Ix'iliiif?<,   111].'!  I;t|t,  ,1.  li.  l,i|i|iiiiciitt  di. 

1.  uiH,  Tiiouias:      niniciil  Disc.r.lira  ot' tlio  Hi'iirt  licat,  I'.  IS.  I1.i,.|,it,  Ni'w  ■S'l.rk.  ll'l:;. 

Hill.     I.cnrianl;      Tin'     Mr.-liaiiisiii     (if     the     ('ircMilatiiiii     uf     the     Itlooil,     in     Scliiifor'n 

riiy>iiil(ij;y,  ii,   ijilKI.      Vciiii];;  .1.   I'fiitlanil. 
i;:i«kill.    \V.    II.:     Tln>   Clint ract ion    of    Cai.liar    Miisil  ■.    in    Srhiifcr'.s    I'li\siiiliiyy,    ii, 

IIMMI,  ^niint'  •'.  rrnllanil. 
h'lM.k.  XI.:      l-'iirtliiT   .V.lvanii's    in    I'liv  siol.i-v .    I'.Mill.      i:,|.   |,v    I..',. nan!    Hill.    K.    .\rriiil.|, 

Liimliin. 
I'.irtrr,  \V.  T.:      .Viniriran   Trxt   liimk  of   l'li\>iiii|ii.'v,  \V.    li.  Saiimlris  (Vi.,   I'.'liO. 


1*  •ri'riiial   I'aix'isi 


111 


iM.i.Williani,   ,1.    .\.,    ,1    „/.:       Ilr;i;l,    l;il.;.    i\.   ;;ii:;;    il.i.l.,    I'lM.    \,    l.-i.'.:    Hiit.    MimI. 
.Iii|irn;il,    .\cn.,    lull;    \I1    Iniiraat.   CuMfirri.s   .if    .\l,-.liiinr.    I.i.n.lnn.    IHI:;,   .See. 
II,  l'liysi..l(i-y. 
-Hill,  l.i'onar.l,  F.  H.  S.,  il  ,il.:      I'm,-.  K,n.  Si.,-..  li.Hl.  /(.  Ixxviii,  :;ll;   ilii.l..    I'.U.-i.  /;, 
l\.\x\  iii,  ."i(iN  ami  ."ilii. 
Krianu.r,  .1.:     .\ni.  .Imir.  I'liysii.l.,  IIIUI,  xxxix,  |nl ;  j|ii,|.,   I'.il.;,  \1,  si'. 
iDiiwn^,  .\.  \V.:     .\rii.  ,l..i.r.  i'liy.-i.il.,  liur..  xl.  ."iJL'. 
li:iylisH,  \V.  M.:      I'm,,-.   Ki.y.  S.i,-.,   ISHC,  Ixxxix,  //.  .'.vc. 
'  KiM.wlti.n,  r.  r.:     .li.iir.  I'liysiol.,  I'.Ml,  xliii,  Jl'.i. 
^Milniy,  T.  II.:     .li.nr.  I'liysi"..!..  1<I17,  li,  J.^ili. 

-i:y,tir  ;inil   .Mi'.k:      |l.;irl,   llMl,  v,   11!P;   il,i.|.,   I'.U,  v,    l:;?;   .\  i,i.  .I..iir.   I'liy^inl.,   lull, 
xxxi\,  liiis. 

■I'lirt.r,  W.  T.:      .\rt.  ..a  <  iniikitiiin   in  ;in   .\iniiiian  TiMl k  i.t'   I'liv-inl.icv.   \V.   H. 

SaumliTM  (■".,   llMiil. 

'   ltr-...lii-,  T.  (i.:      rn..-.  I'hysi.il.  S llMi.-,,  .I,,,ii.  I'liy^ii.l.,   I'.m.-,,  xxxii. 

'iStcuarl,  (1.   .\.:      Heart.   l!ll  I,  iii,  :i.;. 

!-(^;(rriy,  W.:     .\in.  .I.iur.  I'liysidl.,  IMl',  xxx,  l.'il. 

'■Min.'s,  (!.  K.:      .Imir.   I'li\si'.,l.,    lui:;,  \hi.    |ss. 

"iiilm,  A.   K.:      .I.mr.    i;x|.ir.   .M.-.l.,   1!)|L',   x\i,   T.'.L';   l.'.iliinsi.n.  (1.  I'.nil.v:      Ilii.l..   llU:,, 

xvii,  i-2'.<:  Ciiliii  ami  law  is,  T.:     Il.i.l.,   IHi::,  x\iii.  7:;;i. 
1  Malhison.  (i.  ('.:     .I..nr.  I'liysii.l.,  llild,  xli,   I  H'.. 

■  I'.iit.r.  W.  T.:      .\ni.  .I,.nr.   I'liysi. .1.,  Hill,  xx\ii,  L'Tti;  il.i.l.,   1!>I.-.,  wwi.   lis. 
■■.\l:irtin,   i:.  (i.,  an. I  .-..  wi.ik.Ts.         ■i\.  .Ion.-.    I'liysii.l.,    liiM,   xxxii,  ::1l';    \\\i\,   Ji'ii; 

im.'i,  xxxviii,  SIS;    lllli'.,  xl.    . 
"Hayliss,  \V.  .M.:     I'r...-.  I.'oy.  s... ..   i:his,  1\xx,  B.  .;:;'.i. 
Hill.  I..'onar,l:      Tli.'  I'l.ysi..l,.uy   an.l    I>a1li..|..^v   ,.f   ih,.  ( ■.■r.l.i-al   rir.iil;i!i,.ii,  .1.  arnl 

.\.  rimr.liill.  i->;<r.. 

Hill,  I...  an.l  .Ma.'l I,  .1.  .1.  i;.:     .Ii.nr.  I'liysii.l..  lliiiii,  xx\i,  .",!M, 

-iM:i.l.'...l.  an.l  rear..'.  U.  {'■.:     .\in.  .lonr.   I'liysii.l.,  lull,  xxx\.  ■>:. 
r.'ilir,  \V.  T.:     Am.  .I..nr.   I'l.ysiol.,   Is'is,   i.   144. 
Hill.   1,..  ami   Uarnar.l,  II.:      .Ii.nr.   I'liysii.l.,    IssT,   xxi.   .".J.!. 
"  :irl.T,  K.  I'.:     .Innr.   I.al..  an.l  t'lin.   Mel.,   I'.Mii,  i,  71it. 

Kllint  Sniilh,  (;.,  an.l   Tear,  T.   II.:      Sh.ll   Slio.-k.   l...n-in;nis,  (m..  ii   A.   i  ,,.,    p.i;. 
I'.irlcr,  \V.  T.:     Am.  .Ii.nr.  I'liysi..!.,  1liii7.  xx,  :W. 

■>.  .-lij:.  .\I.  (i..  anil  .Iosr|.|i.  I).  I;.:     .I.,nr.  I.ali.  an.l  Clin.   M..I,,   lim;.  i,  ■Js,',;  ;,]*..  s.m- 
li','  ;iii.l   l.yon.  K.   I'.:     Snr;;.,  ilyn..'..  an.l  Ol.si..    I'.ili..  ii.   1 1.;, 

■  " l.rsiin.    Y:in.lcll:    Am.    .Ii.nr.    I'liysiul..    I'.mn.    x\i.    1."..'.:    al.-,.    Mam.:      Hull.    .I.,hns 

II. .|. kins   llos)...   ll'll,  |..   Uld;    .\iark«.-il.|,  .1..  iin.l   .starlin;;.   Iv    I'.:   .1,  iir.    i'livsi.il.. 
I'.M'..   xlvii.   J7.">. 
•M..rr,si,n,   1;.   A..  :in.l    lli...kiT.   I).    1,'.:      .\tn.  .Ii.ur.   I'liysii.l..    l!i|."..  xxx\  ii.   m'.. 
''!l>i.   I'.   II..  SliH;irl.  (i.    N..   :in.l   diilliri.'.  < '.  ('.:      .Imir.    KxjUT.    Mnl..    I'.Mis.    x.    l;ili; 

.-•.•  aisi.   Di.Ilry,  1».  II.:      .Ii.iir.   Mnl.    Iii'si-;i nil.   I'.Mi'.i.  |..  '.).",,  ami    I'.iln,   |..  i;.;!. 
'  iii'»:iy,  II.  H..  ami  .la.ksi.n.   II.  1'.:      I'r...\  Soi'.   Kxp.T.   Hiol.  :in.l   .M.il..    HH."..  xii. 
I'.''!:  KrlanjtiT.  .1.:     «i,.s<-ll,  (iasxer,  I'ro.-.  Am.  I'livsiol.  Soo.,  Am.  .lour.   I'livsiol.. 
i:il>>,  \lv. 


M 


!■      i 
it 


2i).S 


iMi'    riiti'i  i,\ri()N    III'    'iiii:    i;i.uiiii 


■III  ii.li  r^cii,   N  ..  :iii.|    li.-iy^'iinl,  W.   II.:      .Imir.    I'.inl.  rinin..    |!r|s.  NXNJii,  .■'.:'.;i,  ll  l."i  :;:,- 
.■;i;."i  iL:i\rs  (.hlrr   irfcT.  nc.- i.     S,.'  :ils,i  M.-ic  lio.l.  .1.  .1.   i;.:     .I.iiir.  I.iili.  iiikI  <liii 
\\ii\..  (.■•lili.rial  I,   I'.ils,  iii. 
■  ><h..rl.  l.'.iMlrl:     l.iiiir.t,  I.I. 11, 1. .11.  I'.H  I,  |.,  l:;l. 
i\l:iiiii:     .luiir.    Am.    Mr. I.   A-mi.,    I'.pIs.  I\\.  r.ll.      Al>"    Ui.Mi.ii    Mnl.  :in.|   Sihl;.  .Ii.iii\ 

l!ii;. 
■■l':iiii ,  \V.  I!.:      r;i|.,is  l.y  t'Hiiii..ii  ^iii.l  Ci.lhil.i.iiiii.is  ill  .i..iir.  .\rii.  \li.l.  A>^ii.,   I'.ii- 

i\\,  .•.-II.  .■.ji;.  :•■■■].  r.ll.  i;i^. 

■■riirlir,  K.  1..:      I'l...-.  .\iii.  I'liv-ii.l.  S..r,.  Am.  .Imir.   I'li.v-inl.,   llMi;.  xlii,  I'.m;. 
■\Vi;:i.'iis  <■.   ■!..   :iii.l    l>«iii;.    \.    I..:      .Vm.   .Ii.iir.    I'liv>i..l.,    llilH,    xlii,    ITi;;    Am.   .Ii.in 
.\l..|.  s,-..    I'.HT.  ilii,  (■.(•,(•,. 


1* 


i   : 


4 

;1 


¥^ 


"3 
3 


'.-1 


PAKT  IV 

THK   KKSI'lKATIorN' 

CIIAITIOU  XXXIV 
l.'KSI'IKATION' 

Fur  ciiinciiifuci'.  tlic  pliv  Niiilii>;y  of  ic>.|iii-;itiiiii  iiuiv  lii'  I'diisidcrt'd  nil 
■V  its  nil  clKiiiirs,  its  riiiitiiil,  ami   its  i  In  mislrii. 


f 


THE  MECHANICS  OF  RESPIRATION 

nf  tlic  iiiaiiv  I'iiftnis  ciiiici'ImumI  in  inaiiilainiii'^  tlic  iHniiial  fiiiictiiuiiMff 
"f  llir  animal  liody.  tlic  icspiiatni y  ai-t  is  prulialily  tln'  must  impdrtant. 
I'll  this  ai'Cdunt  ami  also  lii'causi'  we  arc  cdnsi-idiis  df  tin'  rcspiratdry 
innvfini'iits,  till'  ]i|iysidld<ry  nl'  icspiiatidn  lias  liccn  stu(li«'(l  I'rdin  tlic 
i.iilifst  times.  .Miicli  dt'  tin-  i-ailitr  wnrk  natiiially  cdnrcint'd  itself 
V,  itli  tlie  study  dt'  the  air  that  enters  anil  leaxes  the  Innirs  at  eaeli  resi)i- 
'.itiiiii  the  Ni'Mtilation  dl'  the  liiiiy:s,  as  it  may  lie  ealled.  Two  dlivioiis 
iiniperties  dt'  the  i'es|)ired  air  are:   (\)    its  piessiire  and   ('1^    its  vnhinie. 

The  Pressure  of  the  Air  in  the  Respiratory  Passages — the  Pulmonary 
or  Intrapulmonic  Pressure 

This  is  readilx  measured  hy  insertiiiL.'  a  tiilie  intu  une  nnstiil  and  enn- 
iii'lin^'    the    tulie    with    a    manometir:    at    eaeh    nuriaal    iiispiratinn    the 

iii.iiiiinieter  reiiistei's  a  neuatixe  jiressui I*  2  nr  :>  mm.  Iljr.  and  af  eaidi 

'  \piiatidn,    a    i)dsili\('    jiressure    n)'    almiit    the    same    de>_'ree.      AlthdU^'h 

'iniin:illy   1)1"  small    magnitude,   the    int rapiilmnnie    |iressui'e   may   lieenme 

11  I  \    '_rieat    when  an.v  (ilislruetinii   is  dtTeicd  to  the   free   ]>assai,'e  nf  the 

ir      The  urealest   pussilile  expiiatiiry  ))ressure  can  lie  measured  hy  sim- 

'i.\    Miiwiiiir  into  a   mercury  maiinmeter.  when    it   will   lie  ('<|Ual   tn  that 

liii-li  all  the  muscles  nl'  the  tlmrax  and  alidnnwu  can  exert  in  enmiircss- 

i'.r  the  luMLTs.     In  a  strduu'  man   it   may  aiiniunt  In  miu'c  than  100  iimi. 

Ml'       Siiiiilaily,    flic    greatest    piissjiiie    iicLraliM'    pressui'c    (in    iiispiratidii 

ly  lie   measured    liy    attemptinj;   td    inspire    tlirousrli    a    fulio   cdiinccted 

'•    ill  a  iiiaiiiimcter.     It   represents  the  t'nrce  with  which  the  inusculatiire 

I.",'!! 


;i  m 


I  ^ 


:t{io 


Tin:  Hi:si-iHATi()\ 


(if  the  lliiiiii\  iiiiil  alxliiini'ii  ran  opcii  up  tlic  tluiracif.  vn^f,  and  im,i\ 
(•<|Ual  -Til  HUM.  Ilir.  TIh'm-  iiH'aMiicmciits  in  tlimisi'lvcs  art'  imt  of  mm  I 
inipiM'taiicc.  cxi-.'pt  as  a  iticasiuc  nf  mnsciitar  (Icvclupini'iit. 

Iiitrapiilniiiiiic   jirisMircs   ilial    aic   iiilfii liatc   ln'tut'cn   tlic   twn   i'\ 

trt'iiii's  will  he  acipiiicd  in  tlic  lower  aii'  passairi's  in  cases  in  wliicli  tlni. 
is  partial  olist  fuetion  of  the  nppei-  respirntor-y  passajjes.  as  in  In'oneliitis 
spasm  of  tiie  jilottis.  diplitlieiia,  etc.  Dniiiiir  i-iiinihiiui  also,  tin-  intiM 
|iulmoiiie  pressure  may  lieeonie  very  liitrli.  In  tliis  aef  tlie  thorax  is  (irvi 
tilleil  wiih  air  liy  a  deep  inspiration;  the  glottis  is  then  closed,  and  .1 
forced  expiration  is  made.     Whi 


lien  a  snttunently  liii,'li  nitrapnlmonie  pre> 
sure  is  attained,  the  irli-ttis  opens  and  the  smlden  chaime  in  pri'ssur'' 
causes  so  forcilile  a  Iilast  of  ail'  that  the  ofTendini;  I'orei^jn  sid)stani'e  i- 
frecpientlv  carried  with  it  out  of  the  air  )iassay;es.  It  is  often  assumi'l 
that  durinu'  eouirhini:'  the  sudden  incri'ase  in  pressure  in  tlu'  aheoli  \\\\\ 
teixl  to  cause  their  walls  to  rupture.  This,  liowever,  is  not  the  casi 
Tin'  aheoli  .jo  not   alone  sur>P'"t   the   inci-ease  of  )>ressurc;  they   UH'rel\ 

act  as  tin'  inner  layii'    oT    a    prai-tieally    Ininnijic ns    structnri in 

posed  of  lunj.',  |.lenra  and  thm-ai-ic  caL'c.  When  the  tissues  of  the  Inn.- 
an-  partially  dei;en<'iat.'d  or  atrophied,  as  in  oh!  people,  then  it  is  po> 
siMe  that  a  rupture  may  take  place,  hut  under  ordinary  conditions  ii 
is  not  likelv  to  oci'ur. 


("     ■ 


•I 


'!     4 


Amount  of  Air  in  the  Lungs 

]\reasnrements  of  the  nnniunt  of  respired  air  liave  recently  assumed  .-i 
(•(Uisideralile  interest  oii  account  of  the  \arious  aiiplicati<nis  which  can 
he  iiuide  of  tlieni  iii  the  stiid.\  of  Inusr  conditions.  Tlie  liilnl  air  is  lliat 
Avhii'li  enters  ami  leaves  the  liiiiu's  with  ea.di  respiration  (aliout  ')()()  cc.'  ; 
the  CDiniilniii  iildl  ulr  is  that  which  we  can  take  in  over  ami  ahove  ai 
ordinary  tidal  lopiratioii  >  nlioiil  I.Miii  e.e.  1  :  and  the  s.ipith  )iii  nlul  nir. 
is  tliat  which  We  can  ■:i\c  out  after  an  ordinar.v  tiilal  expiration  lahoii' 
I'llin  e.e.'.  Takiiii:  tliese  three  tojii'ther,  we  Inive  what  is  known  as  tl:. 
vilnJ  fii/Kii  ill/.  It  is  Usually  ahout  .'i-'ilMI  e.i-..  and  is  i-epi-esented  li.\  tl' 
aiiM^unt  of  air  which  we  can  expel  from  tin-  Iuiil's  after  as  deep  an  iiispi 

ration   as  iiossihle.     The   \ital   i'apaiit>    is  dindnishi'd    ii rtaili   inilm - 

nary   diseases   (see   paire  :;il,.      .Vfii-r  all   the  supiilemental   air  lias  ]«•< 

expelled,  there  si  ill    remains   ill   the   Iuiilts  a   lai'ire   \olui if  air   whi'-: 

can  not  lie  \-oluntaril.\    expelled.     This  is  known  as  the  rtsiihinl  nir.     '{'■ 

iin'asure  it   in  a  dead  animal   it    is  1 essar.v  to  clamp  the  trachea,  open 

the  thorax,  renin-,  e  the  luni:s  to  a  vessel  of  water,  aUil  then  allow  the  ai. 
to  collect  from  the  opened  trachea  in  an  inverted  graduated  cylimh  ■ 
Ono  jiart  of  the  residtnd  air  is  sometimes  called  the  miuimnl  nir:  it   ; 


HI  SI'IKATUIS- 


301 


ii'|irfspiil('(l   l)v   tli.'il    wliicli   is   lint   cNpcllcil   t'roiii   tlic   liiiiu's  of  a  <leail 
.niininl  wlitMi  tlio  thorax  is  niiciii'il.     In  tlio  ciillapNc  of  tlic  lunp;s  thus 

|iio(liict'(l,  the  alveoli  arc  not  coiniilclcly  eniptit'il  ot'  iiir.  1 a\is('  sonif 

lucoiiirs  pocketed  within  them  aiwl  is  expelled  only  when  the  lunits  are 
roinpressed  under  water. 

Tiic  volume  of  the  residual  air  can  readily  he  measured  ihirimj  life 
]>\  cansiiiir  a  i)i'rs(in.  after  a  forced  explication,  to  take  two  or  three 
l.ieaths  in  and  out  of  a  riihher  lias;  eontaininj;  a  measured  ipiaiitity  (d' 
;iii  iiidilTerent  tras  such  as  hydiiiireii.  Siipiiose  the  liai.'  t<»  contain  at 
the  start  4000  e.e.  of  hyilroL'cii,  and  aflcr  a  feu  lircaths  ilOOO  c.c.  of 
tliis  !.MS  and  lOOIj  c.c.  of  othci'  cases  itlie  total  volume  of  hydrofjen  and 
lApireit  air  in  the  liasr  lieiiif;  still  40(1(1  e.e..;  then  the   residual   air  will 


Uutimuui  iii!t|)iratiflii- 


I  vinj't^mfnfal  an  .-^ 


'  TliijlUr>    iu-|rir;ll|. 

TIDAL  AIR 
I  T'lii.ury  i'\[jiratn'ri 


Suf>t>l'm^htl  (111  - 
Ifaximuni  cxi'irUMn  - 


:"\ 


2000  120    ■ 

r  <■        cuh  in 


1500       100 

•or    , 
c.c-     cutiit 


I 


—J  \  Vi?k!  ,;ij,;t,  ity 

jocu&in:     I 


e.tIM('-;cy  tif  (fiuilit.riiim 


.\l!l"tllll» 


111. unci    t'V    tilt"    ];lm'>    ip    vaii'ii-    l>li:l-t  >    >■(    Mtilin.iiv    ;i;i.l    .it     tiirnd 
r«-'pir,ili"n,      i  I'l  ..rii    U';illi  i ,  i 


'ft  *n 


r; 


'■'I-  \'.IV\  e.e.,  for  it  is  evident  that  after  a  feu  hreaths  the  composition  id" 
ilic  exiiired  air  in  the  liatr  will  he  the  same  as  that  in  tile  lunfjs.  This 
lalciilation  is  liased  upon  the  assniii|ition  that  no  hydioiren  is  alisorlied 
'•)  the  hlo<id  duriiiil  tlie  expeiiment,  which  is  not  sirictl.v  the  ea.se. 
ilie  amount  ahsmlied  is.  hnwever.  so  small  in  two  or  three  hreaths  as  to 
inake  it  i)ermissilile  to  (lisiei.'ard  it.  The  measurement  eau  also  he  made 
iy  takiiift  a  few  hreaths  in  and  out   of  a  hai;  coii1ainiii<;  pure  Oj.     B.v 

■  -•I'Mtainin!,'   the   jtroiiortion   of   nitroiren   that    collects   in   the   Iiafj,   the 
;iiaiitity  of  residual  air  can  he  calculated.     AVe  shall  see  later  that  the 

■  i-asuiciticnt  tif  the  residua!  air  duriiifr  life  has  some  practical  impor- 
'■ince  in  eonnection  with  the  ineasuremeiit  of  the  hloodllow  throui;li  the 


■.m 


Tin:    III  SPIUATKIN 


f" 


'i 


Alveolar  and  Dead  Space  Air 

In  !icMitiiin  to  tlioc  iipiicti.'s  i<(  riN|iiri.l  ,iir.  uc  li.ivc  to  (■oiiNiilcr  tl 
.li\isinii   of   111.'   iiir    in    till-   Inii-.'s   into   wluit    is   .'iill.-.l   ,ilrn,l„r   air   .iih! 

iliiiil  sii,ii>    ;iir.     Till'  foiirKr  i-  tin'  air  wliid incs  ii ntact   nitli  tin 

I'pitii.'liniii   tliroiiiili   wtiicli   L-as  ili'Viision   Ih^Iuccii   iIt  I,|oo<1  and   llic  ai^ 
oci'Mi's,  till'  latter  lirinif  tin'  air  \\lii.'li  tills  tlic  ri's|.iiatoiv  |mssa!.'''s.     Tli' 
<l'M.l  s|.a.M.  ,-an  not   !><'  i|,'|in.'<|  anat.inii.-ally  uilli  rxai-titn.li' ;  it   is  fun. 
tiiiiial    I'atlici-   tlian    inni|,hiilHiiic. 

M.'aMiiinn'ht  ,<(  ilir  \<,1mi f  ll„'  ahc..|;ir  :in,l  ilca.l  Npaci'  air  ran  1- 

'"•"'''  ill  ii'i  .iiiiiiial  Invatliin-j  nii.l.r  noriinil  roinlitions  li.\  takin.,'  a. I 
vanta'j.'  ..I'  tli.'  lai-t  tliat.  wliilr  it  is  in  tin'  liinu's.  tin-  air  lias  a.M.'.l  |. 
it  nt,  i.a>.  \\liicli  is  piTsriit    in  111.'  ins|.in'.I  Mi,-  ,,nlv  in   n.'i-liirilplc  tracT. 

''''!'•  " -^ii'y  'I''';!  ■•II'':  '  I      til.'  \..liiiii.'  ..I'  th.'  ti.lal   rcs|.irati..n:  i '_>  ■    tli. 

I"'' "'■I-''  "''  •'•'    ill  ah.'olai-  air:  C]     tl,,'  [..•r.'.nta-.'  of  ( 'I  >    in  tli..  ti.l.V 

air.     Sn|.|„,s,.  tl,,.   xalii.s  to  1...  .'.(10  .•.,•..  (I  ]„.,■  .•.■nt   aii.I    »   p.'r  .•.■iit,   iv 

sp.'.'ii\,.|\  ;   til. Ml   111,.   v,.liinn'  of  alvrolar  air  iniisl    I.,'  .'.(1(1     "^       :'.:;:!   ,. ,. . 

an.l  III.'  il.a.l  spa.-c  Hi?  .■  .•  Tlic  m.^asiiriiiii'iit  so  ina.l.'  is  a.MMirat.'  .ml' 
uli.'ii    ...rlain    piv-'aiitions   air    lal;..n.      M...-aiiM'    .,f   tli,.    i.ra.-li.'al    iiiip-r 

'•■I" ''■  "li^    l''ii'    "'■  "III'   siil,.j,,-t    u,.   sliiijl,    l„,,\,.v,.r,   .l.'f.T   its    fiirtli,  i 

.•,nisi,|,.iati,.n    until    «,•   Iniv.-   I mi.'    fainiliar   uilli    tl,,.   i.r,.n..ial    f.-atiir.- 

of  |.illiM..iiai,v  pliysi,,|o.j\.  Sill.-.,  til.,  first  ail-  t..  imo\.'  into  lli,.  ah.'.ili 
at  III..  l»..jiniiiii..'  of  inspiiati..ii  is  that  pivscnt  in  tli..  .I,.a.l  spa.-.',  tli. 
l.-ist  air  .■Np,-ll.d  fi.nii  tli..  alvf.ili  .,n  tli..  pr..\  i..ns  .■xpiration.  it  i.  ,,i 
n.i  \;'lu,.  in  piirilViim  tli,.  air  aliva.l.v  pr..s,.nt  in  tli.'  ah.'oli.  If  w-  t,,!,,. 
a  ti.lal  iiispirati.iii  as  aiin.iinl  iii..'  t,,  .".(lU  ,. ...  .-m,!  th..  fnn..ti..iial  .l.-a.]  spa,- 

iis    ir.ll   ....v.    if    is   plain    tlial    o,:l>    :;.-(l   ....     ,,f   H ,|,,i,|,.    ,.,1,.    ,_,.||„^    ,|^,. 

"'^'■"'i-  ••iii'l  "I'l'  'li''  siil,s,.,,ii,nt  ,.Npirati,,n  is  .-.iinpos...!  of  i:,il  ,•  ,..  ,,. 
oiltsi.l,.  air  tliat   lia.l  I, ,.!-.■,!  in  il,,.  ,!,.„, |  spa,-,,  plus  V.r.U  ,..,..  of  ,-,|v,.,,|,.|,.  ^iji- 

Tlics..    f;i,.|s  .l.-s,.r\,.   ;i    ..,.|t,-iin   an nl    ,,f   cnipliasis   hc-aiis..   of   tli,.;. 

pra.-li.-al    iiiiporlai in    many    |ili,.|,oni,-na    .-,,111 t,.,|    witli    i-,.spirati,.i- 

' s'I'loiii   tliinks,  f,,r  ..xainpl...  tliat    ,,iit   of  tl,,.  :,{)()  ,..,-.  „f  air  inspire! 

uitli   .-a.-li    l.r.-alli.   only    :;.-,(!  ,■  ,..    r,-a.-li..s   tl,,.   alv.-..li,   vvli.'r..   it    .•.,111, -s   v 
.'onta.-t  «itli  tl„.  •_'.-,(Mi:i(»Mi  ......  „f  „j,.  ,.,i,.,.a.Iy  pr.-s.'nt  in  tliis  part  ..f  il.^ 

Iiiiifrs. 

Til. -I-.,  niiist  tli..r<.f.,r..  I,.,  a  s.irt  of  i,il,  rfn,-,  s.iiii..«  li.-rc  in  tin-  .-iK,...' 
li,'tw....|i  til,,  fr.-sl,  ..nisi.l,.  air  that  .-onn-s  in  uitli  .-a.-li  l,r..alli  tlin.mj; 
tin-  I,r,,h,-!,iM!,-.s  ;,,it!  tli.-  air  wl,i.-i,  is  mi,,i-..  uv  |..ss  sia.jnanl  in  tli.-  ,-il\,-nli 
This  iiit..rfa.-..  must  mov..  l^a.-kuai-.l  an.l  f.,r\var.l  s..in..«hat  with  .-.h-I 
lir.'alh,  an.]   a    rapi.l   .lill'iision   .>f  ..xy._r,.n   ami   of  CO,   must    taU..   i.l.i.  ■ 


ItlM'lHATluN' 


:{(t:i 


J 


.iiiiiss  it  lii'twri'ii  till'  itiMpiicil  nil'  iiinl  tliiit  in  tin'  iilvi-nli.     It  is  iiii|MiH>il)li> 

In  (i\   iiiiy  aiuitniiiic   |iiiiiit   at    wliirli    tlic   intcrtarc ins. 

'I'lii'  iilioM'  )l('siM'il>t'<l  iiii'cliiinisiii   t"iii'  the   .  I'litiliitiiin  <if  tin"  alveoli   iii- 

.  ins  tlif  inaintt'iianrc  ul'  sliirlit  Init  I'lmstaiit  cliaiiurs  in  tli ■iii))iisitii>M 

'if  till-  air  iii'xt  tlif  alvctilar  ciiitlii'liiiin.     It  liclps  tn  pii'vciit  siiiliji'ri  vaiia- 

'iniis  ill  tlic  aiiiiiiiMt  of  irascs  in  llir  lil I,  ])arti<'iilai'l\    »(  CO^,.     Slinulil 

vii'li  vai'ialiniis  (ii'i'iir,  inrunhii'  stiiiinlaliiin  nl'  the  ii'spiraliiry  and  iitlicr 
iiii|"iitaiit  cciitcrs  that  arc  iiillmnrid  liy  tin-  aiiinunt  iif  tliis  iras  jjri-scnt 
ill  simple  Niiliitiiin  in  llic  lilniiil,  wniilil  lie  llic  lesiilt.  Thr  iin rti<iii!sin 
V,  I  ri  >■  lis  II  aorl  of  nil  rlnniiiiil  hiilj'ir  lifi  iHiiniiinliitni  tin  sinlili  n  vhilliijis 
Ml  (/((v  rum  I  iilrnl  inn  iirmliiii  il  hii  iiisjiiinl  imi  mnl  i  i  jiiriil  imi. 

Respiratory  Tracings 

Till'  iiicasiiri'iiii'iits  111"  air  I'cir  tin-  lic'ti'iiiiinatiiin  nl'  tlir  I'mrL'iiiiiLr  Nal- 
!i'>  ail'  liiatle  1>\  tlit"  use  iit'  iiii'lfis  ul'  Naiiinis  l\pi's.  Smiirt  inirs,  linw- 
'  \v\\   it   is  iii'ci'ssary   tn  nlttain   an    iiisi-rilinl   r nil  nl"  tlic   rcspii  atimis. 


I  ills  may  lie  citlicr  ipialitatiM'  or  i|iiaiitilati\  c.      A   iiihilitnlirt    n innl  is 
Ivi'ii  li.\    attacliintr  sonic  suit  of  rcccixiny:  taniliinir  to  tlic  tlmracic  Avail 
■'!•  licst  t\pc  is  sliow  II  in  l''iv'    \^^^>.  anil  conncclini;  iliis  witli  a   ncnrd- 
:    laiiilioiii'   arranircil    to    write    on    a    liiackciicil    siiifaec.      Winn    it    is 
^ircil  merely  to  count   tlie  respirations  or  to  oliscrve  tlicir  i("_'iilaiily, 
'  li  a  tracin;;  is  all  that   is  rei|iiiii'il.  Imt   ohvioiisly  it   ilocs  not   tell   us 
'    »i/(i7i   iiir  has  ciitereil   ami   left    the   luii'^s   at    cacli    resi)iiatiiili.      To 
■liii  a  iimnililiillri   Iniciiii/,  we  iiiiist  either  eonnecl  a  iceoriiiii!,'  instru- 
ct   with    the    trachea    or    inclose    the    Imdy    of    the    animal    in    what    is 
"\m   as  a  lioily    plethysinin^raph.      In   oliseiv  atioiis  on    lalioiatory   an- 
ils the  licst  type  of  r<'ei)r'ilinir  iii^trninciit  to  connect  with  the  rcspira- 
passaircs  is  the  (I'ad  •<'■  I'^rojili  pnciimo'.'raph.     A  ImmIx  plcthysmoi;raph 
i^cil  in  the  case  of  man   is  shown    in    Fiir.    lOM.      All   these   instruments 
-'  of  course  he  oalilirated,  which   is  done  liy  ponriiiir  a  definite  niim 


ri;.  il^ 


•     ■ 


I  'iSk  I  i 


:t(i4 


Till    i<i>i'ii<niii\ 


f     I 


li.T  lit'  <•  c.  lit'  wiiti'i'  I'lmii  .1  u'riiilmiti'  into  a  luittli'  willi  wliii-li  tlic  rn-onl 

illL'  instnillli'tlt    JH  Ciinili'.'tiil  li.\    tllliilllf.      Tln'  ilisplarrni.'iit   nl'  tllr   \MiliM'. 

|iiiiiit  i.'i\i'<i  lis  till"  iii'iM'ssaiy  ilata  I'm'  >tanilai'ili/.atiiiii. 
The  Intrapleural  Preuure 

The  air  \\]i\<\\  ui-  lia\f  Jusl  ln'cii  ciiiisiili'iiiiu'  ili'pi-nils  for  its  mo\.- 
iiii'iit  ill  ami  uiii  lit'  till-  air  |ias>ia>,'i's  ii|iiiii  i-liaii«;i's  iii-ciiriiiii;  mi  the  (iiitri 
nspri-t  lit"  till'  llln^r^  in  the  -ii.ii'r  lii-twcMi  iIhmii  ami  tin-  tlioracic  wall 
This  is  caliiil  tlic  intrapii'liial  spare  It  ilms  iml  really  exist  as  an 
actual  spaee  in  the  lixiiiu  animal.  I'nr  the  visceral  pleura  which  eovei- 
the  liiM'.'s  is  in  accurate  ami  intimate  appiisitimi  with  the  parietal  pleura 
III!  the  inner  asp.  .-t  nl'  the  tlimax. 


i 

1         1 

i  ■  ii 

f 

\\  ihc  tliiiiacic  walls  arc  puncturcil  in  a  liviinr  animal  or  in  one  whii! 
has    recently    ilii  ,1,    the   air    will    riisli    into    the    tlmrax.    the    two    layci 
III'  pleura  separate,  ami  the  luiiirs  collapse,  eaiisiiii;  teniporarily  a  spa. 
tu   he   fiirmeil   hftuecn  the  twii   layers  nl"  iilcuia   ami   iiulicatiiif;   that 
eeitain    siilialiinisphcric   or   iicy-atiM'    pressure    must    exist    in    the    inta. 

thorax  to  iirevent  the  lunjis  t'roiii  colla]isiii'_'.     The  (lei.'ri C  tills  nci;, 

live  pressure  iiia\-  he  measured  liy  eiiniieetiiii;  a  tiilic  ami  a  inaiioiiicl. 
with  the  thoracic-  cavity.  While  tlio  tlioiax  is  a!  rest,  as  in  cxpirali' 
or  immediately  after  deatli.  this  jircssurc  aiiiciiiits  to  ahout  ."»  mil! 
meters.*  ( iii  inspiration  if  increases  to  In  milliniel-rs.  Thero  are  tlici 
fore  two  )>rolilems  to  lie  considered:  (  li  the  cause  of  the  negative  pi. 
sure  ill  the  <|uiesceiit  thorax,  and  rl''  the  cause  of  the  iiicroasc  of  l! 
negative  pressure  diiriiis.'  iiis|>iration. 

•Till-  liiiiiiis  MKM  1,1. lu  alts  lli.ll  Ih,-  1,,,-iMiip  is  nritutiv,.  ,.r  *i.li.itm()-|.li,iii       Ii   i<  a  Mi,-li..n  i.i,,-. 


'■'  -IIHATIiiV 


rwr. 


The  Permanent  Negative  .         ve.     I.i'i   us  stini    uith   th.'  iliain:<> 

Ui.ll  iirCllf  ill  IIm-  lIlnlilN  \\llcn  tin-  lilsl  lili'illll  i-.  i|la\MI  Wllili'  till'  illl 
ilnal  i"  still  III  nil  III,  till-  llllius  ccilillilrlrly  lill  ijii'  lll>iia\.  WIkMI  tin- 
liist  lif'alli  is  iliiivui  tlir  tlu.i  iii'if  I'Miri'  r\|iaiiils  iiimi'  i|iiiclJ_v  tiiaii  lli«' 
liihUs.  Ml  tllllt  Illl'  latter  lirrmiir  st  I  c  I  i-linj,  til.'  sinh'liinu  \\iv,-f  luiliu' 
till-  air  tllllt  in  illl  linjiiricl  ilitn  tlirill  t'luin  the  iiiilsiilc  tliinil'^li  (in'  t  la 
.  I  •  ,1  ami  Iiiiiiii'liial  tiilns  On  liiNMiiniiiif  strelcliicl  tin'  lunuN  fill  tlir 
1  .iiasi'ii  s|>aiM'  iTrati'ii  in  tlir  tliiiia\  lp\  tin'  mralii  r\|«an-iMn  ,,r  tiir 
ihaacif  iMi:f.  This  in  itsi'lr.  |miv\  ex  ci'.  wmiM  iiirt  i'\|ilain  tlic  raiiM-  >>'i  a 
viiiialMiiis|ilicii('  pirssiiii'  ill  tlic  iiitiaplriiial  s|iacc  .\ ii'M iui'  ijilnr  niiist 
..iiiif  intii   play     naiiirly.   tlic  riastic   tissue  dl'  tlic   luni;^.    vvliidi    li\    tin' 

•  \|Mlisiii||  will  Im'I'iiIiii'  strrli-lic'ij  ainl,  tlM'ic  T'li  .■,  IimhI  I'l.n^taiillv  t.i  Ir 
!;;\  til  its  pri'viiiiis  ccpnilitinn  ami  so  cmtI  a  pull  lui  \\\f  si  r-ndui  is  !"■ 
''I'll   it   anil   tlir  timrarii'  \vall.      It    is  tliis  ila^tir   iii'ml    w  liii'li   w,-   nallx 

iii'asuiT    wIk'Ii    \\ Ill I    a    iiiann|ii''li'i'    \'illi    llir    iiit  I  .ipliii!  a!    spa'-'' 

Ihi  ian;liiiiif   lift'  till'  iilliu's  ri'iiiain   nl'  stiiali''i    »i/''   lliaii   llir   tliMia'-ic   uall. 

•:.|    llirl-rriil'i'    111    lill     tlic    tliularir    ra\il>      llii'\     aii ii~t:iiill\      \u<i\<'    <>r 

'' "    ilisli'Milril    illiil    tlir    flaslii'    tissiii'    soiuii'  lial     -I  n  N-liiil        TIh'     Iiiu'.'s 
■■<.   liii\M'\  IT.   not   till'  only    st  lui-luii'v   in    \\\v    lli..i,i\    wliii-li    lir.'iiiin'    i'\ 
iMii'liil:   all    tliin  « allril    xcssrl-   alul    vi^i-i'ia.    lil-i'    tlir    N'in-..    lln-   i^nplia 
ji>.  till'  aufirli's,  etc..   niiist    aUu  ln'i'i'ini'  opmril    mit    a    littl'' 

Wlirii  till'  tliKi-ai-ii-  wall   is  puni'tilii'il  aiul   tin'  mit-iilr  air  allnw'i!    I'l  rr 
i',li-\    In  till'   inl  faplriiral   span'.  ililTiTi'iiri -.   in    pir^^iii-i'    im   li.ii'j.r   .'Ni^t 

■  1'     till'   iiiniT  anil   niitiT  aspi'cts   of  tlir   luip.'s,   sn   that    thiy   i-'illap-i'    inln 

■  •  ■■  piistiiiortcm  ('iiliiiilinii  nil  ari'iuint  o|'  tin'  rlastii-  ii'inil       li'a  pnnitnii' 
■!    till'    thnfarif    wall    ul'    a    lisinir   animal    is    iinnn  ihalil.v     ni-rluihil.    tin' 

:':'.'s  will  cNpaiiil  a'.;aiii.  hi'cailsc  tin'  lilnnij  ahsorlis  thf  -^asi's  I'mni  tin' 
tiaplciiial  spjii't'  fiiiil  f<'iTi'ati's  the  partial  \ai'uuiii  ri'i|iiiri'.|  t.i  i'\pani! 
■  Imiv's.  Tliis  alisiirptinii  of  lms  in  tin'  piriiral  i-a\it\  i^  ii-uallv  i|Uiti' 
I'iil;  hut  if  thf  init  iitiiiitliornr,  as  tin-  cniiilitinii  is  ralliil.  i>  allnwiij  in 
^ist   fur  )iii\    li'n<;tli   of  tiiiio,   llu'  luii-.'s  will    imt    lii'i-niiir    prnp<'rty   i'\ 

'h'll  au'aiii. 
The  Greater  Negative  Pressure  on  Inspiration.     Tin'  i'a\  it\  nt'  tlm  tlm 

lirrniiii's  iuiTcasiil  ill  all  iliaiiirti'i's  iluiiinu  in^|iiral  inn.  with  thr  fi'- 
'  'hat  a  jrri'ati'f  space  in  the  pleural  ea\il>  has  tn  lie  lillnl.  All  the 
''  waned  strnetlires  in  the  thorax  theret'ole  h iiie  vtill   innri'  ^tret.hed, 

luiii;s  of  coiii'se   partieipatiiiir  to  the  '.'featesl    i'\triii    lin-au-i'   n|'  the 

•  '.inee   of  outside   air.      Tlie   stretehill','   ot'   the    elastie   st  lllet  11  res    eaUs.'S 
-leater  pidl,  of  I'        lti\e  ])resslire,  to   he  eMlteij    ill   the    pleural    e,i\ity. 

'lad  of  lieiiit:     •'>  niiii.  llir,  as  in  eNpiiatioii,  the  iiitral  Imraeie  piessnie 
^    I'liines  to  he  alio\  e     Id  iiini.   \\<j. 
N\  hen  any  ohstructinn  exists  in  the  air  passa'j:es.  Ihe  eh.'iu'jes  in   intr;i- 


H'W: 


I  If 


'MW, 


Tin:  i!i;si'ii{ATi(i\ 


f 


i  t 


V.' 


l-^i 


I 


if.' 


^1  b^ 


tliiii-iU'ic    lucNMiic    iHcMliii-fil    l(\    the    incivcinciits    iif   rcspiiiitimi    liccoiii 

iiicm'  i>riiiii,iiiirr(l   ili.iii  umlci'  iriiil  ciiiiilitioiis.     Wln'ii  tlic  thorax  ex 

1>;miiIs  with  the  t  imcIh'.i  lihic^kcil,  the  lliiiys  aii-  ii.it  aide  tci  iiju'ii  up  siitti 
'■i''ii'l\  '"  till  all  thr  ^pa(•(■  s..  that  Ihcii'  is  cxfi'ssivc  (liJatatinn  of  th. 
vein-.,  aiiii.-hs  aiicl  .■Miphai.'Us,  as  well  as  ijiawiiii:  in  <.f  the  iiitcicosl,! 
spac-cs  ami  liiilu'iiiir  iipwanis  dl'  the  iliaphrauiri.  IT  a  iiiamiiiictcr  is  cdn 
iii'ctcij    with    tin'    pliiiral    spai'i'    iiihIci-    tJH'si'    i-iin<litiiins,    a    vcrv    lar-^i 

iii"_'ati\  1'  111'  Niicliiin    picsMiic   will   I Iiscivi'il.  aiiiiiiintiiiit  (il'tcii   In     7i 

"!■  ^"  'ii'ii  II'-'-  ll  i-^  piissililc  tlial  miller  siicli  I'oinlitiniis  sniuc  spar. 
iiiiLiht  tciiipuiaiily  cNist  lictwi'iii  the  parietal  and  viseeial  laveis  cif  th- 
pleiiia.  lint  it  eiiiihl  nut  remain  hni!.'.  fur  it  wnnld  ve?-y  snoii  lie  lilje'l 
ii\  tliiid  esndinir  linm  the  lihind  vessels.  In  the  nppnsite  ediiditidii,  in 
x>.  liieh  \hr  respiiatiii-.v  passaues  are  lili.idved  and  a  rorecd  exiiiratimi  i- 
iiiaile,  as  I'cir  e\aiiiple  in  the  first  sta^'e  nl'  eiiii",diiii;,'  nr  diirin«;  siieli  aei 
as  defeeatiiin  and  part iiritiiiii,  the  thmaeie  eaiic  is  cdniiu'essed  ii|)iiii  tin 
viseeia.  with  the  lesnlt  tliat  the  air  in  llie  liiii^s  assiinies  a  jicisitix. 
pressure,  ainoniilinir  nlti'ii  t.i  nearly  1(1(1  nun.  Iltr.  lr  a  iniiietiire  wniinl 
is  inaile   in  the  Ihnrav   under  thes.'  I'niidil  imis,  the  lung's  instead  of  enl 

lapsing'    will    liiilu'e    out    of   tlie    wniiiid.    I'.ir    what    is    reallv    Mirrini;    i- 

that    the  lliorax   is  f.ireilily  ediitraetiii^r  nil  dcelnded  sacs  d|'  air. 

It  is  the  alternaliny:  ehan;.'es  in  intiapleiii  al  pressure  that  are  resp.in 
silih'  fni  the  ehanues  in  iiitrapiiliiidiiii-  pressure  anil  these  i'lir  the  iiinv.' 
"I'll'  "I'  'lii'  ill  and  nut  of  the  Iniiirs  with  eaeli  respiratimi.  In  dtlic 
wi.rds,  the  thiiraN  dues  iint  expand  on  iiispiralidii  lieeanse  air  riisli, - 
i'l-  !i-^  ""'  uninitiated  iiiiaj^ine.  hut  air  rushes  in  liecailse  the  tlnnax 
iNpands. 

The  Influence  of  Intrapletiral  Pressure  on  the  Blood  Pressure.    'Ih. 

ind\einents  nf  r.spiration    pnuliiee  etVeets   mi    the    vaseiilar   system    tlia 

■I'"'    "I    ' sidiraliie    iiupiiitanee    in    iiiaiiitaininy:    the    eireiilatiini    nl"    th 

hliiiid.      It'   an    arterial    lihuid  pressure    traeinjr    is    cxaiiiineij.    it    will    h 

iilisiiveii  ilijit  aside  riiiiii  tl ardiai-  pnlsaliiuis  larire  waves  i'\ist  iin  it  tli-i 

are    appiuximalely    s>  nehriiiinns    with    the    respiratory    movements,    tl 
iipsliiike  lit'  eaeh  111'  these  waves  eiiriespoinliiii;  in  i.'eneral   with   insjiii.' 
timi.  and   the  duw  iistiuke   with  expiratiim   (  l-'iy:.  2'2i.     These   lespiratni 
variations    in    lilond    pressure    iiiii;ht    lie    due   either   to   ehainres    in    Ilea' 
rhythm     or    to    a     purely    merhaniral     eanse.      Ke^'ardin-.'   the   (irsl    pus- 

liility,  it   is  indeed  tl asi'  in  most  animals  that   the  pulse  is  ipiieker  <■ 

inspiration  than   on   expiration,   hut    that    this  alone  is   not    an   adeipnit 
explanation  of  the   rise   is  shown   hy   the    faet    that    it    still    persists   »([■ 
the   vavMis  eonlriij    of  the   heart    has   lieen   eliminated,   either  liy  enlli' 
the   nei'M-  <,!•  hy    t!--^'   aetioi!   iif  atrtipitif. 

The  eanse  must   therefore  lie  a  nieehanieal  one.     r.eariiit;  in  mind  li 


■  -^ 


i    1 


I   1 


UKSI'IHATIOX 


no? 


I  iTccIs  wliifli  wo  liii\('  sci'ii  iiic  produced  mi  llic  inovciin'iil  (if  nir  in  iiiid 
out  of  the  Illiijjs  Iiy  the  cli.iiiu'cs  in  <-!ipiicily  of  tin-  tlior;ix  \\illi  f.icli  its- 
Iiir.ition,  wv  iiiifniMlly  assmiic  tlnit  the  inci'ciisc  in  lilood  jirossurc  nniy 
lie  dnt'  1()  till"  fiK't  tliiit  on  inspiiMtion  more  Idood  is  .sucked  out  of  tlie 
-.\steiiiic  veins  into  those  of  tlie  tlioiiix,  llint  this  excess  wlieii  it  is  i»ro- 
l>.llcd  I)y  tlie  licMit  into  tlie  arteries  raises  the  li!o<id  pressure,  and  that 
nil  expiration  the  ojiposite  eonilitioii  olitains.  That  the  nio\ciiieiits  of 
llie  thorax  on  inspiration  do  accelerate  tin'  sjieed  with  whieli  the  venous 

111 I   is  traveliiiir  towards  the  heart   can   easily   he  shown   hy   measure- 

iiiciits  of  Iiloodllow. 

This  exjilanation,  liowever.  does  not  snlHii-e  to  aeeount  for  iill  tho 
iliaiiiies  of  lilood  pressure  which  occur  in  ri'spiratiou.  lor  if  we  take 
\(iy  ae<'urate  tracimrs  of  lilood  ]>ressure  and  of  the  respii'atoi-y  iiiove- 
iiiciits  si<le  liy  side,  we  shall  find  that,  allhouyh.  in  ",'eneral,  the  lilood 
pressure  rises  with  inspiration,  yet  the  liey;iiiiiinLr  of  the  rise  is  eonsid- 
I  i;ilily  delayed-  that  is,  ininiedi.ately  follow  inir  the  Iie<jiniiiii<r  of  the 
Inspiratory  aiM  the  arterial  lilood  lu'essure  continues  for  some  time  to 
fall,  c'lnd  at  llie  liejriiinin;;  of  exi)iiation  it  continues  for  some  time  to 
li^c  (|"i'_'  'J'iK  JIoreiiM'r,  it  will  he  found,  if  traciny:s  taken  from  dif- 
rerent  animals  are  compared,  that  freipiently  tlie  iiciieral  effect  of  ex- 
piration is  to  ciiuse  more  rise  than  fall,  and  of  iiis]>iraf ion  more  fall 
Ih.iii  rise.  It  will  lie  found  that  these  differciKM-s  are  dependent  larirely 
nil   the  type  of  respiration,  whether  thoraeie  or  jilidominal    fl.ewis')." 

l.el  us  eoiisid"r  first  of  all  exactly  what  will  happen  in  an  animal 
hrealhiiiir  entirely  liy  the  tlinni.r  (e.  jr.,  tlie  ndiliit).  The  first  efl'eet  of 
the  inspiration  is  to  (■aus<'  tlie  veins  leailinir  to  the  auricles,  the  .lurieles 
tlicinselves  iind  the  Mood  vessels  of  the  llinirs  fo  lieeomo  suddenly  ex- 
panded. More  lilood  theii'foie  will  How  into  tliem.  l-'or  a  moment  or 
tv.o  this  Mood  will,  howexcr.  tend  to  stjimiate  in  the  moi<'  capjieious 
\ '■^sel^,  :iiid  it  will  111-  some  time  until  it  finds  its  wa.\'  to  the  left  side 
'il'  Ihe  heart:  therefore  the  initial  elTect   of  ins|)iratioM   is  a   ilistiiict    fall 

in  aiterial  lil 1  pressure.     When   the  extra  spa -realed   in   the  lil 1 

\'NN(|s  h;is  lieen  lilled  with  lilood,  th.it  is.  when  inspiralion  has  pr,n'- 
'iiiillv  ccjiscd.  fill'  Mood  will  llow  on  in  increased  volume  to  the  left 
-iile  of  the  heart,  and,  therefore,  raise  the  arterial  hlood  ])ressure.  On 
•  rjiirnlidu  tlie  lii-st  effect  is  that  the  diminishiiiii:  ne-rative  pressiii-e  will 
'Miise  the  tliin-w  ,illed  vessels  meiitioiieil  al'ove  to  constrict  aiiil  thus 
-'|iiee/e  the  Mood  inside  them  into  the  left  side  of  the  heart  and  I'aise 
ilic  pressure;  liut  the  ultimate  I'lTect  in  the  liiter  sla'_'es  of  expiration 
'hII  III-  that  the  vessels,  lieinj;  eonstricted,  will  allow  less  lilood  throUi;li 
:'"'II1  and   the  arterial   Mooil   pressUle   will    fail. 


Tal. 


ike  now  the  ease  of  itftilintiiinil  n sftinilimi.     In   inspir.-ition   the  di: 


if) 


;l 


m 


U 


l-i 


k 


\   lit! 
I   ii|j 


\i  H  ! 


:!i)S 


nil     lilsl'IU  \Tln\ 


Uf 


i 

=1    ■> 


|ilil;i2lii    (I.'sitikIs    mikI    i-io\\,U   III,.    \iM-ii;i    iii,'iiiiist    llir    vena    cava,    willi 

''i''  i«-iilt   tlial   at    tii-s|   iiKiiT  111 I   Is  s,|ii,>.vc.l  into  llio  flinrnx  anrl   lli. 

''I '    l'i'"->uic    l>'ii(ls    sli-liil\     I..    iIm..      Afl.T    this    initial    ctTc'i,    li,,« 

fM'f.   tlic  (-(.Miiirosion   «{  till"   vi'iia  cava   ranscs  less   IiIikkI  to   vcnrh   tin' 
tliiiiax,   ami   the   aitciial    lilooil   iiicssnic    t'alU.     Tlic   condilinns   will    hr 
i\ai'tl>    rcM'iM'iI    on    i'\|)iiatiiin.      Tlii'   initial    clTcct    of  tlioracic    iiispiia 
tioii   is.  tlii'iit'iiri'.  til  iiiaki-  ttic  arterial   liloml   picssiirc   fall,  and  tlic  in 
itial    I'ff.'cl    (if  alMlnniinal    iiis|iirati.,n.    (u    make    it    rise.      The    net    ('(I'l'i-l 


■    <    '    .    ^    •    ■. 

4    ARDOMCN 

/..^ 

('/> 

B    CHCST 

t.p 

■       - 

BPi.n,                 J ' • 

Intp^^ 

z    i-P__ 



Intp 

• 

Chest 

D  cues T 

f.^ 

"^ 

-—^ 

,nsp 

Im.  IM.  lITi.t  i.f  ,il..l..inHi.il  .111.1  ili,,l  lir.iithiiiR  nn  tlir  piiN,-  .in.l  lil..o,l  |.r.'«sMr.-  of  iiMi 
Al..lnniiTial  in-|ii.i:'nii  r.ii^.  -  i!m-  |.r, -mim.  .,,„|  ,|jmi„i,.l:,.^  ,|„.  .nriii.lilu.lp  .if  llir  ,.iil«v  rurvi-.  TiM'aM, 
iii>i.ir.iti.,n    !,-,    .I,.ii:.     I,,«,r,    t!..-    i.n -.ur.-        Iai  i!:ili..n    ha-,    il„-    .^pprvit,-    rftViX.       l  lM..m    l.rwi-- 

produced   will   be   tlic   atu'cliraic   sum   of  tlicsc   two   opposing,'   influences 
(see  Fitr.  110 >. 

Another  factor  that   comes  into  pla.v   in  determiiiiiiji  tlie  effect  of  tlie 
respiratory  movements  on  the  cardiac  output  acts  throus;li  the  chanKes 
in  the  pericardial   inessure.     \Vlien  tiiis  is  lowered,  as  carl.v  in  iiispira 
lion,    it    encouia-res   diastole,   thus   eausinir    lietter   fillin';   and    thei'efore 
lietter  discliari;e  from  the  liearl. 

These  considerations  tahen  tou'ctlier  nial<e  it  easy  to  understand  the 
cliansres  in  lilood  pressure,  jiarticularly  in  tlie  veins,  which  occur  when 
.'1  forced  iiis|)irator.\  or  expiratory  tiiovenient  is  made  witli  tlie  sflotti- 
elosed.      .1    fiirriil   <  ijiirntiiin   of  this   natnr ciirs   diirini;  the   acts   oi 


1 


El 


KKSI'IHATION 


;!()!» 


.Ict'ccjiliiHi  iiii(i  piirtiifitioii.  iis  well  as  in  Iho  (ii-ht  stiincs  of  ciiuj^liiiijj ;  it 
is  iilso  |>n((liic.'(i  liy  lilnuiiijr  into  a  tube,  <ir  a^'aiii>t  siniic  rcsistaiici'. 
•  Ill  afcdiint  <))■  the  iiositivc  idcssurc  that  is  linmjrht  to  licai-  on  the  veins 
iis  they  enter  the  tlioiax.  the  venons  piessuie  stuldenly  lises,  slowins; 
iloun  tlie  lh)n-  of  hhxxl  thiousili  tlie  eapilhiiies  and  eausin<;  Iml^inf;  of 
the  veins  and,  if  tlie  elVect  is  siist:iineil,  eyaiiosis.  On  tlie  arterial 
side  of  the  vasenlar  system,  after  a  nioinentary  rise  eaiised  liy  the 
s(|iiee/.in<.'  out  into  the  left  side  of  the  heart  of  tlie  hlood  in  the  eapil- 
l.iries  of  the  lnii!.'s,  there  is  a  more  ])eriiianent  fall  in  i)ressiiie  duo  to 
the  fact  that  less  hlood  is  now  ^''ttinji  finni  the  rii.dit  side  to  the  left 
side  of  the  heart.  After  some  time  the  juessure  lieirins  to  rise  a^ain. 
partly  on  aeeoimt  of  the  liaek  pressure  through  the  eapiilary  vessels 
and  partly  l)eeause  of  \  asoeonst  lielion  as  a  i-esull  of  asphvxial 
I'oiiditioiis. 

Ill  the  opposite  condition,  duiiiiu'  n  fnrcul  in^jilnil'irii  iiiari  mi  iil  ^\  ith 
the  L'lottis  elosed  or  with  the  iiiouth  attached  t,.  s,,iii,.  tiihe  throU!,'h 
"hieh   till-  attempt    is   made   to   siiek   air.   the    ihmaeie   eavities   ojien    U]) 

^^ilhou1    the    Inn-rs    heinir   ahle    t -iipy    eomplc.tely    the    extra    space. 

Till'  dilatation  of  the  veins  and  other  thin  nailed  stiuctures  in  the  tlio- 
iii\  thus  causes  nu  immeiliale  fall  in  lioth  the  venous  and  the  arterial 
I'l.ssiire  ill  the  venous,  liecaus.'  th.'  hlood  is  sucked  tou.ird  the  larire 
vessels  in  the  thorax  and  lunirs.  and  in  the  arteri.il.  hceause  the  Mood  is 
now  delay<'d  in  its  jiassai/e  from  the  riuht  to  the  hd't  side  <d"  the  heart. 
If  this  eoiidition  is  maintained,  the  arteiial  presMir,.  uiay  recover  .soine- 
\\hat.  hut  that   in  the  \eins  is  i>ermanently  lowered. 


t 


■ 

Hh 

V 

in 

Rlt 

1 

•    «i 


Id 


'I 

% 


VI 


11 


■i-J\ 


*\ 


I    Ml 


ft  I ' 


ij 


CllAITKi;  xxxv 
Till-:  .MiiciiAMcs  <ti'  i{i:siMi{.\Ti(t\  ^(^.lltM) 

VARIATIONS  IN  THE  DEAD  SPACE,  THE  RESIDUAL  AIR  AND 
MID-CAPACITY,    AND    THE    VITAL    CAPACITY    IN    VARI- 
OUS   PHYSIOLOGIC    AND    PATHOLOGIC    CONDITIONS 

llv   1{.  (i.   I'l  Aii.  i;,   ll.A..  Ml). 

Dead  Space 

I'liili  T  iiii|iii;irv  cipinlitiiiiis  ol'  Infiilliili'^  tlic  dfiid  spjicf  is  fiiil'ly  cnii 
nIiiiiI    ill   Miliiiiic.      Iliilil.'tiic     iiiid   lli'iKliTsmr    liclicvc  liint    it    may  l)i'   iii 
crcascil   liy  4(M)  pci'  fciit    in   iiiaxiinal   ilccp   iH'catliint;.   and   tlial    the   in 
iTfasc  is  liui'  to  llu'  passive  stn'tcliiiii;  of  liu"  lnwcr  air  sacs.     Altlmu;:,'!! 
sui'li  iai-ii'  variations  in  tlii-  capacity  of  the  dead  spai-c  lias  not  liccii  oli 
served   liy   Krnjrli   and   i/nidiiard'   or  liy    If.   (I.   rcarcc,'"  it    is  uiidoui)li'l 
that   iMiMlcialc  rliylliiiiic  variations  may  occur.     Kvcn   in  dccpir'  Ideal li 
inu'  I  l.'idd  e.e.  or  ovcri,  a  sliulit   increase,  wliicli  witii   ma.ximum  li'vatli^ 
may   .imontit    to    lUO  e.e.,  can   lie  demonstrated.     Tliis   is   not   surprisini; 
wlieii   we  reineiiilier  tiiat    tlie   wall.,  of  tlic   lironelii  and  l»roncliioles  ari' 
made  up  largely  of  readily  (xpansihle  tissue  (elastic  aiul  siiiootli-muscli 
liliersi.     As  the  respirations  liceomc  deeper  i:n<l  tlie  cxpandinj;  forct>  oi 
tlie  inspiratory  movements  of  ilie  tlioi'ax  jpccomes  more  jn-onouiieed,  tin 
diameter  of  the  iiinnehi  and  Inoiieliioles  will   erdarije   iiroportionately 
I'.a!    is.  the  diameter  or  i-ireiniiference  will   inerease  ni  direct   i)ro|iortioii 
to  this   force;  liul    the  aica   of  the  er'oss  section   of  the  liroiichi    (i.e.,   th- 
lapacitv  I   will   increase  as  the  sipiaie  of  the  diameter.     This  depends  on 
the   fact   that   the  area  of  a   circle   is  increased  hy    Ti'i  per  ei'llt   when   lie 
diannlei-   is   ineicasecl    hy   .")((   per   cent,   and    hy   ahout    o()(l   per   cent    win' 
the  diaiih'ter  is  increased    \i\    |(l(l  per  ce?it. 

Th''    capacity    of    the    ijc-ad    spac'    has    :i    I'erlain    eliniciil    si,_r|ii(icai 

Sieliech''  has  estimated  tha.  the  deail  space  may  ini-iease  hy  lOO  ee  in 
asthma,  lint  others  helieve  that  the  increase  may  lie  jireater.  One  re.i 
son  for  the  disi-niil;iiit  residts  lies  in  tlie  fai'l  thai  the  perceiitaire  "i 
( "n^  found  in  the  alveolar  air  olitaine<l  liy  the  llaldane  Priestley  iiicth'"! 
has  Iieen   used   as  one  of  the  hasic   (iirures   in   the  determination   of  tl' 

;{io 


■1 

I  -I 


im 


-M 


tin;    mm  IIANKS    (IK    Hl;sl'IU.\TIU\ 


311 


.•.i|i;iiilv  111'  the  iiir  |iii.ss;ij.n's.  As  cxiil.iiiiril  ilscwlicrc  I  |i.ii!i'  :!4-H.  Ilic  |M'i>- 
luii^Mtioii  of  CNiiiratinii  i'i-i|iiii'i'il  to  nliliiiii  tin-  siiinpli'  of  iilvcoliir  air  liy  this 
hii'llioil  <;iv('s  li<;iii'('s  tliiit  :ir(-  too  lii<.'li  i-vcn  iindi'i'  iioiinal  coiKlilioiis. 
,ini|  it  is  pljiin  tliat  tliis  cncpf  A\ill  lie  <'Najj['_'('i'at('(l  in  astlmia,  wlicrc  tin; 
i\Iiiia1ioii  is  "ircatlv  pi'oloiiiicil.  An  iiK-i-casc  in  tlic  capni'ity  of  tlio 
.liail  spare  mnst   lie  a<'roni])anir<l  liy  an  increase  in  the  respiratory  vol- 

! if  tlie  alveoli  are  to  Ite  a(le(|natel>-  ventilated.     It   lias  lieen  tliousflit 

liy  siiine  clinieiaiis  that  the  dillienlty  in  asthma,  emphysema  and  ear- 
ili.'ii-  decompensation  may  lie  in  part  in  an  inci'eas(i  in  the  dead  space. 
I'.iirliil  estimations  of  the  dead  space-  in  these  eoinlitions,  however, 
r.iil  to  deimnistrate  any  fjreat   variation. 

.\ii  explainition  of  the  fact  that  the  di'ad  space  in  eniph.x  sematons 
pilicnts  has  heeii  fonnd  to  he  >;encra!ly  lai'yc  when  determined  by  the 
li;iM,jne-l'rie.stle\  method  (see  pa^'e  :14l().  anil  alsn  I'cir  some  of  the  clin- 
ii  .il  phenomena  accompanyinu'  the  eoiiditjun.  nniy  he  ns  follows:  In 
rinpliysenia  the  walls  of  the  ahcoli.  especiall\-  alioiil  the  lateral  and 
liiucr  horders  of  the  lunirs,  lia\f  lost  their  elasticitv  and  fail  to  exjiand 
'If  relax  )>roperly  ;!'  .inir  the  respiratoiy  c\ele.  As  a  restdt  the  air  in 
Ihise  alveoli  icniains  relatively  un.  iianu'ed  exi-ept  when  forced  i'es])ir'a- 
limis  ai'e  made.  When  a  sample  of  alveolar  air  is  taki'n  directl.v,  this 
iliad  air  is  i»ushed  ont  of  the  distemled  and  diseased  ahcoli  hy  the 
t'lifced  respiration  rcipiired  in  the  direct  samjilinsr  of  th(>  alveolar  ail". 
>iiicc  the  air'  in  these  ahcoli  has  lieen  in  contact  with  the  hlood  enter- 
iii'.;-  the  liinsis.  it  has  a  hi'_'li  CO.  i-ontent,  which  I'csiilts.  when  comiiared 
\'illi  the  nnifornd\-  low  ("(>_.  content  fo\ind  in  the  tidal  nil',  in  udvin'r  a 
!  live  Iji^nre  for  thi'  dead  space.  Since  tin'  caiiacity  of  the  dead  space 
i^  imt  incr-iNiscd,  the  lilood  in  the  inirmal  alve<ili  is  prohalilx-  lieinir  sujier- 
Miitilatcd  in  oriler  to  comix'nsate  foi-  the  hiv'h  ('<V.  tension  in  the  hlood 
■  iitciinu'  the  left  heart  from  tlie  diseased  alveoli.  However,  the  <\, 
I'liitcnt  of  the  hlood  leavin>r  the  sonnd  alveuli  is  |)ractically  normal  (Ite- 

MiM'  supc  rvenlilation  can  not  canse  it  In  take  \ip  moreV  ami  can  not 
■ 'iMii.ensate  for  the  low  O,  content  in  the  Mood  coniiinr  fi'om  the  dis- 
'  i^i'd  ahcoli.  tlie  net  effect  heiiii:  thei-efore  a  low  tension  of  0_,  in  the 
'I'liiid  leaxiiiir  the  heart,  which  acconnis  for  the  cyanosis  oft(>n  seen  in 
1  iMIihyseiiia  d'earce'l,  A  somewhat  similar  cxpliiiiation  can  he  triven 
i'"r  the  cyanosis  ])resent  in  ]>\ilmotiar>-  edema,  if  we  assume  that  all  the 
.il\ro!i  ill  this  condition  do  not   share  alike  in   the  edema   (Hoover). 

The  Residual  Air  and  Mid-capacity  of  the  Lungs 


I'uriiii;  muscular  exercise  the   residual   air  of  the 


Inntrs   is  ini'rcasi'i 


the  vital  i-apacity  decreased  Cr.olir'i.     This  causes  the  hint's  lo  as- 


'   l>l 


I- 


$1 


9 


t  I 


It      ! 


^     ir 


i'i 


I   '■■! 


ir. 


Tin;  Ki;si'in.\'rioN' 


Mllrir  ;i  liiuic  iiilljiliil  ciiiidil  idil  Ixtut'cri  lirciitlis  iir,  jis  it  llllS  Ix'i'll  cllllii 
>il\  vt  \  Iril.  ;[  vi'iilir  mid  ciiiiiifit^ .  Tiii'x'  cliiinu'cs  iiiiiy  servo  as  .i 
|i|i>  >iii|ii'jic   Miclliiiil    inr   ini-ri';i>iii'_'   llic  (rtirifiicy  ot'  iilvciiiai'   v<'iitilati"i 

s"  a--  t ri  till'  iTiiatii-  iii'fils  cii'  till'  l>ii(ly.     This  is  partly  Ipccausc  tli 

puliiKiiKii  >  \i>s>c!s  lifciiiMc  liilatcil  ami  the  IpIooiIiIhw  through  the  iiiiii; 
is  liniHi'd.  and  partly  liccaiiNc  ot'  tlii'  iiitliit'iicc  (d'  tlio  reserve  and  sup 
pliiiniital  ails  on  llic  tcnsiini  ul'  tlie  arterial  liliiiul  jjases  during  the  ic^ 
piratur.v  e\  i-je.  j'oi-  e\ampie.  it'  the  liiiitrs  were  (•(>nii)letely  depletc-i 
i'\'  ail'  diiriiiL'  e\piiati()ii.  the  IiIimmI  lea\  iii<:  them  at  the  end  of  this  ;ir; 
"oiild  li.'  iiiiir.ly  veiioiis.  Oil  tin-  other  hand,  if  tile  amount  of  air  lei' 
ill  the  liiiius  ai  the  eii(|  iil'  expiration  Were  above  tlie  normal  iwaount 
raeji  iiieiciMi-iit  <>\  (  ( »  Liixeii  o(T  from  the  hlood,  or  of  ()^  ahsorhed  li\ 
it   \voiiM  piodiice  Ic^s  cdiaii'.'e  ill  the  ]iress\iie  of  tlie  CO.  or  O.. 

'i'lie    iiiipdrtaiice    of    lliese    iiilludiees   will    he    seen    fi'oni    the    followili',' 
li'.'iiies.     If  the  residual  and  supplemental  air  amounts  to  "JdOO  e.e.,  and 
the   pi'feentaLre  of  ('(1.   in   the  ahcolar  air  at   the  end  of  expiration   is 
.'>  p.T  cut,  till  n  |iHi  ,■  ,•    ,,f  ("u,  must  he  present  in  the  lunfis.     In  a  eon 
(lit  ion  of  lpiidil\    lest   ahoiit   L'l)  e.e.  of  this  <;as  is  excreted  during  a  res 
piialoiy   eyele,  so  that    if  the  hreath   were  held  dnrin<:  this  jK'l'iod,  tie 

pel iil.i'ji'  of  ( '( I    would  rise  fiom  ,">  to  i;  |u>r  eent,  and  an  inspiration  oi 

• would   lie  tf'ipiiied   to  liiinir  the  air  in  the  lunjis  hack  to  '■>  pc^ 

'■'■n'   >'(  <  "        '"11   the  other  hand,    if  the  residual   and  siii)i)leniental   aii 
aiiiriiiiitid  to  .',{»»)  e.e.  with  .">  per  eent  of  CO.,  in  the  alveolar  air  at  tin 
end  ot'  the  expiration,   there   woidil   he    l.'iO  e.e.   of  CO.^   in   the  lini<rs  a' 
the  end  ot'  I'le  expiration,  so  tliat  holilinv  the  hreath  for  the  time  of  tie' 
iespii;i1oiy  .•>  ele  woijhl   raise  the  iiereeiitaire  of  CO_.  only  to  5.66  fpro 
\  ided  the  prodiietion  of  Co.  was  the  same  as  in  the  (irst  easel,  and  a' 
inspiration    of  lllli)  ,-i\    would   he   iiei-essary    to    reduce    it   to   the   norma! 
.'Xpiiatc.iy   tiuuie.      Or.    puttiin.'   it    another  w;iy,    the   ])rodnction    of  CO 
can   he  i, Ideas.'. I  ."id  pcT  cent   in  the  lime  of  a   respirator.\-  cycle  without 
;i(TectiiM_'  til.-  tension   of  liases  in   the   lunjis.   provided  the   residual   an. I 
siippHiiieiital    air   and    tin'    xolunie   <i{   the   resjni'ation   are   inereased   ."id 
pel-  .('lit.     If  ..iii\   one  of  the  f;ietors  is  ehaii<re(l.  liowever,  then  the  Iial 
.iiic''   of   till'    i.'spir.ition    inust    he   dislurhed,   and    the    greater   variatioi' 
in   the  tiiisi.iii  .,|'  ill,,  .jascs  in  ilic  arterial   tilood  must  oeeur  at   the  dil' 
t'.'.nl    ph.is.'N  of  til.'   respiratory  e\  «de.      Hohr  and  Sieheek   have  show  i 
tli.it   til.'  i.'si,|iial  air  is  iii\  ariahl.v    increased  in  emph\senia  and  that  tin 
iiii.l  .-apacit;    .,f    tli,'    liiiM.r^    is    lik.-wise    increased:   and    it    would    ajjjx'a' 
fr.iiii  Siili...-k's  .lata  that  a  simil.-ir  .•.■ndition  must  lie  present  in  eases  c' 

,|:;'.,„:i":|l-::t..|     he;:rf, 

I'alietits    siiiTcriiii.'    from    (|_\spiiea.    particularly    those    sufTcring   fror 


THE    MKrIIANICS   OF   Rt^l'IRATION 


313 


inliac  (lyspiioii,  ciiii  not  lucaflic  as  cinnfortably  whi-ii  lying  as  when 
itinsr.    This  ciUKlitioii  is  known  as  orlliopnra.    The  advantasc  of  the  sit- 


n 


Hi.'  ovt-r  the  lyiiic  position  for  hrcatiiinfj  i-an  not  he  satisfactorily  ex- 
pliiiiit'd.  The  jfi-eatcr  vital  (•ai)acity  in  the  n|)riKht  position;  the  favor- 
ins:  of  the  refuin  of  the  venons  blood  from  the  cerebral  vessels  by 
;,'ravity;  the  increased  caliber  of  the  i)u]nionary  vessels  because  of  the 


nilar^'ed  thoracic  cavity   (see  pajre  :!!S);  arid  the  iiicreas..  in  the  reserve 
nil'  of  the  hniffs     are  all  factors  to  be  considered. 

The  Vital  Capacity.-  At  one  time  it  was  thon^'iit  that  the  vital  capacity 
if  the  Inncs  was  related  to  their  ventilatory  capabilities,  but  for  years 
the  determination  of  this  value  in  patients  has  been  considered  unimpor- 
i^iiit.  Hecently  I'eabody  and  Wentwor4h"'  have  called  attention  to  the 
t;ict  that  patients  with  heart  disease  become  dyspneie'mcne  readily  than 
ijo  healthy  subjects,  and  that  this  tendency  seems  to  dcj^end  largely 
nil  their  inability  to  increase  the  dejjth  of  the  respiration  in  a  normal 
ni.iiiner.  They  find  that  this  inability  to  breathe  deeply  corresponds  to 
.■I  iliiiiinished  vital  capacity  of  the  luiifrs  as  measured  in  a  spirometer, 
li.v  the  volume  of  the  greatest  possible  expiration  after  the  deepest  in- 
-|iiiiition.  They  believe  that  any  condition  which  limits  the  possibility 
Ml"  increasinsf  the  minute  volume  of  air  breathed  must  be  an  important 
i;ii'tor  in  the  production  of  dyspnea. 

Ill  normal  adults  the  folIowin<r  averaircs  (Table  II.  were  secured  from 
.1  l.uRe  series  of  clinical  eases.  The  subjects  are  •irouped  into  two 
•Lisses.  each  proup  beini;  subdivided  according  to  heijiht. 

TAm.r   I 
The  \ttai.  i'apacih   .,p  tmi:  I.i-\c.s  i>r   \,>»mai.   Mai  ts 


III 


NI'MBEB 

STrrnKh 


14 

44 


HEir.uT  IS 
ft.rt  AM> 

i.N-ciirs 


Over    5' 


5'    ,1"    to 
5-  8IJ- 


N'lRMAI. 

VITAL 

CAPACITV 

C.C. 


.'.mil 

4.tlOl) 
4.0IMI 


M 

MBKK 

WITHIN 

10 

'^  OF 

N( 

RMAI. 

0 

-tl 

-U 

IIir.HKsT 

\ITAI, 
CAPACITV 

."^.KIIO 


I  ■IWEST 

VITAL 
CAPACITY 


.'.11.10 
4..1IIII 


Mir.H- 
tsT 


141 
IJl 


90 
S6 


M'MBKR 
BFI  t>W 

90';  (II- 

\"I1\IAI 


The  Vital  1"ap.acitv  of  the    I.im-.s  ok  Nobmai.   Females 


III 

Over   5' 

ft* 

s.m 

5 

4.11/5 

_',«ll(l 

IJ4 

C() 

J 

n 

flvir  .■;' 

4"     to     ■-■ 

.1.II.MI 

Q 

.1,4:.' 

.'.i.f.n 

ii:; 

SS 

2 

:l 

y   4"   or 
trA< 

j..*<-'.^ 

u> 

.'.SJII 

J.'OO 

I.!  5 

.1*9 

' 

<  I'l-ahody   and    Wcntwortli.  I 

It  would  appear  that   in  normal  jieopli'  the  vital  capacity  is  at  least 
~  '  prr  cent,  and  almost   always  •)()  i)er  cent  or  more,  of  the  standard 
i'ljitcd  lor  each  jrroui).     In  elderly  persons  a  slight  decrease  from  these 
'iiidards  may  be  expected. 


^^H     ' 

f      5 


i1 


1  i 


i? 


"■'1 

I 


'»! 


If ' 


111' 


i'rl 


W 


I 


:UI 


Tin:  iiisi'iuAiiiiN 


Tmii.k  II 


Tin     IfiT   \I|M\    (It     rill     \IT\I     t'M'MIIV    (IK   THK    I.IM.S    Ki    I  III     I'|I\!1M.   CoMHIIIIN    IS 
I'MliAT.-l    Willi    IIIIMIT    IHSIVSK 


(■ \IV\( 

riY 

III  i:  (IK 

(     VSl  S 

■I  All  TV 

OK  HI  niM 
ri  NSM'IIIN' 

1 

'.HI     - 

-."> 

'1 

0 

II 

7ii  l( 

'.til 

II 

.'» 

■J 

.SNMI'TOMS  U(i|;K 


:>\ 


III 

IV 


III  to 

I'll. I, 


"1,11  till  ..(-( 
ill  IP'. (I-  (tl,lll  ■■'I 
»lil.li   lliiv    fill 
«ln;lit    IM  rliiil 
M.liiv  mil.  r   p.ili 
111-   li..-i.|l.il. 


l:l  .mai;k.s 


Few  Nvni|itiim«*  ii-t 
cnililc  III  licaif. 

liistiiiy  (if  (l\s|iii(i 
Willi  (■xcrtimi.  M  • 
iililc  Id  (Id  iiiiiiii  I 
lite  Hdik. 

I>>s|i|i('ii  »iMi  iiidil 
('  r  ;i  t  (»      cxcrciM' 

I'i'W   lllilc   Id  «dlk. 

I'iciliiilili'M,  u  i  I  li 
iiiiiikcd  ni;;ns  dl 
(-  :i  r  i]  i  ;i  c  iliMil' 
licicncv. 

{I'c;ili"ily  .Hill  W.'iilwdith.) 

^    wii      ti^ii!    <i\ii,il    tinii^   :iii.l.   i.v\ii.i'    111   .  li  nii.'i  ^   ill    llii-    \il.-|1    I'ai.iiifv    Iliiv   .-ijilii.ii 
i-i.,.il..      In   ill.    ■■.ii.il.ilio"  (..I'liiiii    III.  V   ai.    Ill,  lii.li.l     .nil    in   tin-  I.  ».  -I    Kri.ii|.   ii.l  . 
■  S\int.f.'iii^    ..i    il  .  ..III!-.    isiiIiLii"    lll.ll..ilr    il\s|,fi,  .1    uliitr    .il    l(-t    ill    111. I    ..I    MM   VI  r\ 
Iii.l.  r   "»...l,iiit"   .111-   iiul.i.li.l   ..i.U    Ih.i-i'    ^iitM.ilK    :il    Willi..    .111.1   aMi'   |.i   (..iilin.i. 
i.u  111  (■.r..ni.   il   w.  ri    al.lc  t..  w..rl..  t.ut   llity  ;iri    ii.it   iin  lii.lc.l   as  llii  y   wcic   vtill   in 


III 


r.| 


liiii 


Tiililc  II  slmws  lli;it  lluTc  is  a  rciiniikalily  clusc  i.'latiiHisliip  lictwccii 
tlic  I'liiiii'al  cdinlil  inn  of  canliai"  |>ali('iits,  |>aili('iilarly  as  rcsiards  tin' 
ti'iiddu-y  til  <|\N|iiifa,  and  tlic  vital  capai-ity  cij"  the  limirs.  I'caliiMly  and 
Wfiiluortli  liclicvc  that  tlic  dctciiiiiiiatiiiii  id"  tlic  vital  capacity  affiinN 
a  clinical  test  ;is  Ik  the  i'liiicliniial  ciduiitioii  <d"  the  licart.  since  cunipcii 
Silted  patients  wild  ill)  not  ciiinplain  <d'  dyspnea  on  cxertioii  liavo  a  iim- 
tiial  vital  capacity.  I'atieiits  willi  iiioi'c  serious  disease  in  wliom  dyspnea 
is  a  |ii-oniinent  syiii])toiii.  have  a  low  vital  capacity:  and  the  decrease  in 
\ital  cai)acity  inns  parallel  with  the  clinical  condition.  As  a  jiatieiil 
improves,  his  vital  capacity  tends  to  rise;  as  lie  liccoiiies  worse,  it  leiiiK 
to  fall.  In  oIIk  liseiises  in  which  iiieehanical  conditions  interfere  willi 
llic  niovenieiits  of  the  luii>i:s,  the  tendency  to  dyspnea  corresi)oiids  closel.v 
to  the  ilci-reasc  in  the  vital  <'apaeity.  The  cause  of  the  dccrcjise  in  tin- 
vit.il  capacity  of  the  liiip^  in  canliac  decompensatioii  is  diflicult  to  e\ 
plain  salisfactoiilx .  It  may  lie  the  limitation  in  the  iiioveiiieiifs  of  tli'' 
liliiL's  piiidiieed  liy  enijoiirenieiit  of  the  ]iiiIiiioliary  vessels,  hy  the  we.'ik 
iiess  of  the  intercostal  innsides.  the  riyiidity  of  the  Imny  thorax, 
eiiipliysema,  or  aceiiiiiiilatioii  of  fluid  in  the  pleural  cavities. 

In  cardiac  disease  the  air  in  the  luiijjrs  at  the  end  of  a  uoriiial  expiration 
is  iiMia!!\  iiii-tiMM-d.  TliiN  is  sitiiilar  to  tin-  cnhditinti  wliich  atlctids  ex-'r 
else,  and  is  ]irolialily  a  physioloyie  adaptation  to  ^'ive  o])tiiuiiin  aeration 
to  the  Mood,  as  explained  alio\e. 


CIIAITKI!   XXWI 
TIIK   .Mi;(  HANKS  OK   HKSI'lIfATlOX    (CnntM) 


THE  MECHANISM  LY  WHICH  THE  CHANGES  IN  CAPACITY  OF 
THE  THORAX  AND  LUNGS  ARE  BROUGHT  ABOUT 

l!v   K.  (i.   I'l  Mi.  i;,   1!.A..  Ml). 

Till'  clL-iiiycs  tliiit  t.ikf  pliic't'  in  llic  rnriii  iiiiil  llic  iliiiifiisiniis  nt'  tin' 
llicii;i\  (liliilii.'  i<'s|iii;itii)ii  ;iir  liimi'^lil  jilmiil  liy  iikpv niii'lits  cif  tlic  lilis, 
.ii,i|ilii;ii.'iii,  stci'iuiiii,  iind  vcrti'liiii'.  Tlic  sliiiii'  wli'n-li  t-.u-h  |>l;i\s  iiiiisl 
III'  cipiisiili'icd   st'piUiitcIy. 

The  Movements  of  the  Ribs 

'I'Ih'  liist  scM'M  luiiis  cif  I'ilis  i)i(i<;i-('SNi\  c'lv  iiiciciiM'  in  Icnirtli,  iinil  ai'i' 
iiliu'licd  (lirccliy  In  llic  .sicinnm  Ity  (•;iitilii'jrini)us  liiimls.  Tlic  ciirlitli  In 
llii'  IwcH'tii  ]>iiir.s  |)|•<li,'|•^■^^i\  fly  (Jci'ii'.iM-  in  Icniitli.  iinii  iis  t'.'ir  jis  tlic 
Iriilli  tlicy  iii-c  iiiilii-t'ctl\  ;ilt;icln'(l  In  tlic  stcinnm  li.v  ciir1il;ii;t's  uliicli  join 
llic  scvcMtli.  The  clcvciifli  iiiiil  twclllli  liii\c  tlicir  ;iiitci'ini'  ends  free,  and 
may  lie  ennsidci'cd  a  ))ait  u{  the  alidnniinal  wall  and  not  an  intfinsic  i)ai't 
uf  llic  llHii'acic  cafto. 

Kacli  ])air  at  rilis,  tof^ctlicr  w  itli  its  aitieiilatiiiir  cartilau'c  and  vcitclx'ip. 
t'liriiis  iv  rinjr.  tlic  iilane  ot'  wliicli  is  directed  fcnwai'd  and  dnw  iiwaid. 
Till'  si>inal  artienlatinns  of  the  upiJcr  lihs  dilTer  fioin  tlmse  ol'  tlic  Inwci' 
"lies.  Tn  the  foriiier  the  artienlatinns  nn  the  tillicrcle  exist  as  eniivcx 
nviiid  facets,  A\liieli  (it  into  cnrrcspnndinir  linllnw  facets  on  the  tiansxersc 
I'lnccsscs  of  the  vei'teliia',  while  the  cori'espnndiiii;  facets  of  the  lower 
nlis  are  flat.  Kach  transverse  jirocess  from  aliove  downward  is  tilted  a 
!  nlc  more  hneUward  than  the  one  ahove,  so  that  the  aiijilc  at  which  the 
'lis  arc  set  to  the  sjiinr  increases  from  ahove  downward.  This  uianner 
:'  ;iitici(latioii  of  the  upper  rihs  with  the  \-ertelii-a'  prevents  any  rotation 
i  :  the  spinostornal  axis,  so  that  there  can  he  no  so-called  liilckct-liaiidle 
:  ivemeiit  in  this  reu'ion  (Kcithl.  The  articulation,  however,  allows  the 
■  '  I'k  of  the  I'll)  tn  rotate  in  an  axis  apitroxiiiiatcly  traiis\'erse  to  the  1)od\-. 
I'll'  allele  which  the  shaft  of  the  rili  makes  near  its  neck,  to^'dliei'  with 
i'  ('  ari'h  of  the  shaft,  which  is  directed  dowiiwaid  and  forward,  has  the 
•  Ti'i-t  of  eausin<r  the   trans\erse   lotatioii   of  the   neck   of  the  rih   to  lie 

315 


I    ftl'S 


316 


Tin:    KOIIHATION 


\^ 


c.iiivcrtt'il  iiitii  ill)  ii|i«iii<l  iiiiixciiiiMt,  wliiili  i>  urciil'' .  in  tlint  part  nltl 
sliiii't  l,\  iii<;  |>;ii'iill<'l  ti>  I  III-  iisis  <it'  I'litiitiiiii  III'  till'  iii'i'k  i  h'lir.  111). 

Till-  ii|i|iri'  rili>.  H'liiM  till*  lii'st  III  till'  lit'lli  riii'iii  a  I'Diic-sliajii'il  tii|i  to  ll 
thmax,  wlii'ri'as  tin-  louci'  niii's  fniiii  a  vcitii-al  vrics,  cacli  ln'iiijt  Mluat'i 
aliiiiist   iliri'i-ily  aliiivc  \\^  iiriL'lilmr.     Tlio  ii|i|ici'  set   is  ai'i'aii<;('il   for  ll 
r\|>«li>iii||  III"  till'  riiiiiral   llpjH'l-  liilir  of  tin'   liin)»s,  llic  lowiT   for  tile   i 
patision  of  tlic  iiiiiii'  or  li'ss  r.\  liinlrira*  lowrr  liiln'->.     Diiiiiiit  iiispiratin 
till'  aiitci'iipiistrrinr  iliaini'tiT  of   the   ciiiiical    piii'timi   of   the    lliorax    \r, 
I'lM'asi's.   lii'i-aiiM'   llic   rilis.   Inu't'tlicr   with   tlii'   NtiTiiaJ   coiincctiiins,   iimiv 
tliroiiL'li  priii;ri'Nsi\rl\  iticrt'asini.'  aii-lu's,  ami  racli  lower  rili  triids  to  omi 
riili-  till'  I'ili  just  almvr.     'j'lii'  inaNiMial  iIm'  «(  llic  I'ilis  rroni  the  lii-st  to  || 


.1.    1. 1-1    ,!•. 


.ll      MIt.lil.l.      /', 


:h    cinr-jl    \, 
each    case. 


rt(l>t,i    diul    rill 


i>l'    riitatiMii    kilo 


ti'iitli  (liiriiii;  inspiiatiiiii  sliil'ls  iiinrt'  aiii!  inin'c  fimu  tlic  aiiliTim'  to  \\\ 
lateral  aspi'ct.s  of  the  tlmrax,  lifi-aiisc  tin-  aii'.'lc  foriiii'ii  liy  tlio  shaft  nr-.t 
the  nock  of  the  rili  aiiproailu's  nearer  to  tlie  articulating;  joints  on  tli' 
vcrtdiru'. 

An  examination  of  the  shape  of  the  tiist  ril>.  its  relationship  to  adjaiii 
structures  and   its  nioxeniciits.  shows  that   it   differs  from  the  otliers 
its  lespiratoiy  function.    The  first  ))air  of  rihs  and  tiie  nianid)rium  steii 
are  Imund  closely  to<;ethcr  liy  short,  \\  ide  coktal   cartilajrcs,  and   form 
strnclnral    unit    which    Keith'    calls  the   fhorinii    niit rnihnn.      This  lid    : 
articulated  liehind  n\\h   the  lirsl   tlinracic  \ertclira   li.\-  a  joint,  which   i 

niiii'c  neaily  tians\cisc  than  that  of  llie  rest  of  ih stal  series;  and  ; 

front  Mitli  the  nianulirium,  which  i-i  also  articulntod  witli  the  elavicl' 


^t 


TIM:    MlilliVIrs   iir    HI  "I'lH  \TliiN- 


•111 


i 


il.i.vr  mill  «illi  the  Im.cI,    .,r  till-  >t)'iiiiiiii  IhI.iw.     Thr   t'rfc(l..ni  of  iiiiiM- 
i.iiiit  at  the  ;iiii;li'  wliii-li  tlic  iiiiiiiiiltriiiiii  iiuikcs  witli  the  stiTinini  lit  tliin 
j.illt   i>  Iflali'd  In  tlic  t,\|ir  .i|'  lni'jilliiinj.      When   llir  liiui'l-  |"illiM|i  of  llir 

•■tii'iiiiiii  is  clcviil'd  (liiriiin  iii>'i'ii!ilioii,  tlir  iiio\ciiii'iit  of  till'  joint  is  not 
li.c.  liiit  wlifii  tlif  stri'Miiiii  is  ii'lriii'tril.  the  iiiovi'iiii-iit  at  tin'  niiirlf  iniix' 
■  iHi  'lilt  to  III  .  I,j|i-k  of  iiioM'iiirnt  of  ilic  sici-iiai  iiiiiniilti'iiil  joint  lins 
I'll  I'onsiilcrcil  li\  NoiMi'  iiiiysii'iiins  as  oih'  of  tlic  iirrtlisposiiii;  caiisos  iif 
I'lliiioiiiin  tiiliiiTiilosis  Diiiiiiir  inspiiatioii.  tin'  first  iil>  iind  its  antrrior 
.ittaclmii'iits  arc  raised  l.y  tlie  s(  aleiii,  and  serve  as  a  point  towards  wliieli 

'li"  s d.  tliiiil.  fointh  and  liftli  rilis  are  elexated.     Diirinif  expiratiuii. 

;liiv  are  di'pri'sseil  toward  the  lower  rilis,  whieh  t'orni  a  more  nr  less 
ii\iil  iiasc. 

The  eoiiiliiiied  ctTeel  of  these  inllneiiees  is  to  proilnee  a  motion  of  the 
'i|i|>er  rilis  uliicli  is  deserilied  \>\   the  eliiii<-iaii  as  lieiii<r  ninliilnliirii.     This 

\eiiicnt    is   more  app;irent    in   the   upper   part    of  the   thorax,   heeaiise 

li'  re  the  rehlti\e  dilTerenei'   in   the  jenjith   of  the   rihs  is  u'leatest.      Hoover 

.'iiihiites   a   eerlain    diaynostji'    simiilieanee    to    hiss   of    the    iindidatory 

■  i\eiiieiit.  diiiiiiiiitioii  in  the  exteiisiliilitv  of  the  iiiideri\'iii<;  luiiirs  eaiisint; 

!   Ii.  heeonie  less  or  to  disappear.     The  |ilh     omenoll   is  elieiteil   liy   plaeilli; 

■he  lip  of  the  rinir  tinner  on  tin'  ^ nd  rili  in  the  midelavieiilar  line,  the 

■ip  of  the  middle  liniror  on  tlie  tliiid  lili  midway  lietween  the  mididavieii- 
111-  and  anteroaNillary  line,  and  the  tit>  of  the  iiide\  (iiii;er  on  the  fourth 
m!.  ill  the  midasillary  line  The  pati.nt  is  then  inslriieted  to  make  a 
iii'iclerately  rapid  and  deep  inspiration.  The  tin<!;er  on  the  third  rili  will 
'••■  ohserved  t-i  move  farthi'r  than  that  on  the  seeond  rili,  and  the  fiii>rer 
'  '  the  fonrtli  rih  will  move  farther  than  that  on  the  third.  The  iiiove- 
li.i  III  of  ejiidi  rill  from  alMi\e  d>iwii\\ard  siieeeeds  and  exceeds  that  in 
till-  rill  just  aliove. 

When  there  is  a  nioch'r.ite  dejii'i f  impair;  'eiit   in  the  ventilation  of 

I  •  upper  lolie.  the  three  rilis  move  in  uni.son  a. id  throii<rh  the  same  dis- 
'  ''  ee,  so  that  the  uiidiilatory  inovcincnt  is  lost  altlioimh  the  rilis  involved 
!i!.i>  oxhiliit  n  considcrahle  esciirsion.  The  tnidiilatory  nnivcmcnt  is  also 
I'   paired  liy  any  disease  whieli  ciicroae)  is  on  the  air  spaces,  invades  the 

'eistiiial  tissue  of  the  lunsr.  or  dis|il..,es  the  lunj;  as  in  the  case  of  nn 
'  iartred  heart  or  a  distended  iierieardial  >ae.  AiKither  jxissihlc  factor 
'  tliis  phenomenon  is  that  any  inllainmalorx  jiriiecss  in  the  liiiitf  or  nd- 
.     1  111  tissue  Avill  produce  a  retlex  inhiliition  of  the  muscles  of  the  rihs, 

!  thus  limit  the  expansion  of  the  thorax. 

Tlie  axis  of  movement  of  the  lower  ribs,  as  of  the  upper  rilis,  accurately 


-ponds  with  that  indicateti  iiv  their  articulation  with  the  vertel 


I. 


iiw. 


tl 


ic  muscles  attached  to  them,  as  well  as  the  diaphrafim,  inlluence 


ii-  movements  to  a  larsre  extent.     Anteriorlv  the  lower  rihs  frc 


the 


;t 


.^.iAmS 


TIT 

t 

1 


I  ill 


1 


ill 


31H 


III     Id  >llll\l|ilS' 


!«i\lli  to  Ihc  triilh  ,iiv  jc.i I  111  till'  ^IfiniiMi  lis   llii-  cilitiliii.M'H  H  liicli  inn' 

the  siMli,  M'Xfiilli.  ri'.'lilli.   iiiiilli,  nil. I   Itiilli,  so   lliiit    iiii\    iiiovcinciil    i 
wliirli  till'  lilis  air  niiM'il  is  .ii'i-iiiupjiiiii'il  li\   iiii  jiiiti'iinr  m.ivciiu-iit  iif  ll 
sici  iiiiiii  (  Kiir.  1 12,1.    Till'  I  ills  art'  so  iirtn-iilnlril  In  tin-  spiiiiil  I'ltluiiiit  lli. 
Ilii-  iiisiiiniliiix   lift  i-iiilscs  llif  Ijili'ial  ami  aiiliTim    pail  "I'  cai'li  iili  an  i 
In  iiiiiM-  I'liiwai'il  ami  iiiitwanl  liKHf  tliaii  (lie  uiii'  iilinvt'  it. 

Ill    iialiiial    liiralliiiiy    in    lln'   slaii'liiiii    nr   silliiin    piisliiic    llicic    is 
sliiihl  I  .iti  iisiiiii  (if  III!    >/)//ii    iliiiiiii.'   iiispiralimi.     Tliis  sitvi's  to   iiirriM^' 
all  iliaiiifl.is  III'  llif  tliiira\  ami  its  alisciin'  is  iiiiiliiiilitriil.v  all  iiiipoitai.' 


l-ij.      II   >        l,.,u    1     ImIi'    .1    111.      It    ■'.('      ll-i.'    ll.         'll    .|..T-.il    I.,    ll'.      nil     \.tlilii.i.    sell    fniiii 

li..i,T        .       .It nil     |.i.«.-^;     ./    ./',     ....il.t;     r,    .  ^.-i  hiiMi^ ;     /.    ;.|..ilMir     in     l.ii-|..n    ..»     .ll.ii.Iii  .iki'-i     ' 

l.i.-.i.;.-    ..f    Mill    ..n.l    ilil.li..i;     '.     '.     \    llll..l.,ili     I  s|    ..isi.ii^    ..I     I.  11. till.. M~    i.lil.l     ..I     i|..i|.lll.lk'lii.     ; 

..sl.il     ].,ili..n-.     iiL'lil       111.1     Ull.     "I     .ll  i|.l.l.ii!ll -.  I.  .     '1.     Ilklll     nil-     ■■>     .lln|.|ir.lwlii;     V.     ".     ml' 

nil,  I.  .'-I.il     ii;'i-,l.-,     .vl.i    ll    .-111     ii'.-iiil     11. .11     th.      i.ll.'i.il     ...I ...I-.M      ll     I..11'-     .,    :iil.|    ...     111.      ■ 

I,  rii;il    ml.  ii..-l.il-:     ;    .     '  .,    >i|l,..-l.il    iiiii-.  I.  ~    ..I     lill    -il.         lI'LUn     l.i-.  lik.i   I 

i-iiii1iiliiiliii\  lai-tur  ill  rnliiriiiu  tlii'  vital  i-apai'it\  of  an  imlividiial  «  li' 
lylii!,'  "II  tlic  liarU.  Fiifiiifs  Kivni  liv  lliitcliiiisnii  Tnr  tlic  rlTc-t  wlii' 
liiistiiic  has  nil  till'  vital  raparitv  an'  nl'  iiitvicst  liccaiisc  (if  tiirir  licai  i' 
nil  till- caiisi' 111' 111  tlinpiii'a.  Ill  tin- same  iii<li\  iiliial  lie  rmiiid  tin-  I'lillnwii 
\  ital   rapai'itics: 

Slamliii!,'  4:!lMt  I'.c. 

Sitliiitr  42(MI  re. 

Siipiiic  :!S(I(I   c.c. 

I'lniii'  :i(il>0  va: 


^1 


Till      Mil  II WHS   IIK    KIM'IKV'I'liiN 


The  Action  of  the  Musculature  of  the  Ribs 


aio 


III   il   crIH'lIll    \UI.V,    tl Xtfl-lllll    illlcrciisljil    imisrils   inil>     III Iisiiii'lrii 

,i>  II  lii'iiiiil  fxtt'iiHiiiti  tif  till-  Mfiili'iif  iiiiisflcN  iiM'i'  llir  tliiii'.'ii'ii'  walls,  with 

till'    lilis    as    illtrlNi'i'linlis.       Till'    srillrni    mTM'    In    li\    IIm'    |iii>ilicin    nl"    till' 


B 


I    >W         111  llll -I.ll      Mill.,   I,  s      ,,|        -.h      .111. I      t.lU       «|.|c,^  ■,       .1.1,        %.,„;       ;(.       |,,„k       >,,„;       II  III, 

I  rv,;  \.tiil.t.i:   I.  >iti   111'  .111.1  ..iiiiLiB.  ;   '.   i.   \\    Ih.h.t.-  ,...i.,     ,„,    ■.   j,  ,m.i,i,iI  mt, ,,  ,,.i.,U: 

■f  .     '.    iiililll.il    i,,I,  ,,.i.|.il..    ,  N|,.,.^.i    In     I,  Ml..>.il    ..I     111.      I  M.  ,ii,,l     111,...  I     .         I,,       I      III,,. ,M,iii.,'l 

;      g  111.  r.-..l.iU     1,1     tl,,'     l,,t,  1,  .,,tll,lh:ili'.t,.     .1.,,,,.;     I,,     tt     II,.  ,,      .tl.      II .i.iU     II,,,,      til,       \,It,t.,.il 

?  .     inn. II         ll-l..m     ,\ll,  I,    'I'll.. 111.,. II    I 


i 


i^ 


lii'sl  rill  so  tlial  il  I'liriiis  an  aiirliniiii;!'  I'm'  llic  in-tinii  nf  tlii>  i-xtrinal 
ihliii-iistal  iiilisrirs  ill  r.'iisiii^'  tln'  luwrr  lilis.  '{"Iiry  alsii  laisf  llif  ii|i|)i'r 
ilii'i'i'  |i:iii's  of  rilis  alniit;  with  tlic  iiiaiiiilii'luiii  am!  strriiiini. 

Till'   rillK-liiili   til'  till-   iiil)  iTiisldl   niiisilis  has   liiTri   the   siiIiJimI    ,,(  iiiiii-h 


^N« 


\ — /- 


I  u-      111,       ll.iliih.ri: 
I'.ll    ml,  ti,,.t:il>. 

Wlun    tl,..    ril..    fl,     :,,,.!    ),,/    ,,:,,,    i,,t„ 
1.1'..    I/,/,    ,.    yrt.lUr    Ih.iii    ,i/.i;    tji,-    .1, 


,]it-Mui    t.i    .1,  .ti..ii.t,.il.     111,      tiiiu  tu.n.ii    nil'. I,:. .III. II 


lit,  I  ii;il     .III. I    lA- 


.itMii.    a.l    .-;:;. 


III.      int.  r.i,<::il 


''■  1    lli.in    (..'l;    ll„.    |>I>, 
..     .it    tl,,.    iiit,.iii.,|    ,1,1, 


riniiii    .//    ,iii.v,.    .iw.iy    I't.'in    lli,    %(rU'lii;il    ,'.>ti,itiii    ah    I/'/ 


ot    III,'    ,-\l,ti.i,l    ,,it,-i,,..t.il.    .tl-    •Iti.rltii    I, Ft    ,.    «i,;il,  r    l!,;ii,    .i/';    ;iii.| 
i.t.ll.    .  i-     I,  ,,l;I1,'  „     l.t     i.     t"!,.!!,,     tll;i,i     /.jt.        'I'll,'     r,\'r..'     .u.llt.     ttluil 


|.,. -ill, .11    i.    t.-ik.  .1        III...,,    1.1 


llrmtlii     /'/n. „../,. 


'  1: 


i 


.[■: 


h     I 


I     ! 


i- 


:12() 


Tin:  KK>riRATi()x 


ilcliatc.  and  can  not  hi-  said  to  lie  (h'finitol.'  sottU'd.  Tlio  dii'  ctioii  of  tli 
liltcis  ill  the  inti'inal  intoreostals  indicctcs  tliat  they  arc  exi)irafoiv  ii 
function,  since  tlicy  can  not  shorten  in  tlic  inspiratory  position;  whiji' 
on  tlic  otiii-r  hand,  the  fii)crs  of  the  external  infcrcostals  can  not  shortcii 
in  tlic  ex|)iratory  ))<isition,  and  hence  must  he  considcre<l  inspiratory  ii 
character  (Kid-  ]l-{>-  In  n">l  llanilier>;er  showed  that  nicchanically  tlii^ 
is  the  case,  and  frave  the  schema  shown  in  \'\<z.  114. 

The    function    of    the    iutcrcartilajjinous    muscles,    however,    must    \>'- 
inspiratoiy,  as  is  shown  in  I-'ijf.  11."). 


■:^ 

n^. 

I.ik 

Is 

,.|,„IU 

Th. 

rili~ 

will 

^Ii 

the    an 

1  al 

s 

•  M      Wliu 

which 

haw 

i.l,.- 

.1    III.' 

■  itiK 

>s 

.    ai.'l 

tail 

•..„slv.    ai, 

.i|.|i 

iff 

It   that 

III. 

i.ti 

l\     II,:: 

>.  !u-ir.i  111  .U-tiK.it-tralc  tli.it  tin-  (iiiirM.ni  ..i  tin-  iMtt-riial  iiitcn  ai  liIaKiii'Uis  niliT"  - 
■  al    witii    tliat    iif    tin-    cvtcnial    ititiTiKH-iMu-i    iiitt-ri  Mstals. 

an. I  (I'-tal  lartilaKf  H'^iy  tn"  r<  tanlt-.l  a^  t-mK  lurit  at  llu-  aiiKlf-  n. ./  ami  f>,-f.  r 
pcular  t"'i"ls   c   anil   c   rcim-^cnt    tin-   :>*>  inpluviH   tiiMmcii   tlu-   Imiiy   ami   tin-   tariilaKin"  ■ 

h  trartiiiii  is  n'a.Nv  I  turinK  in-ipirali'Mi  tin-  •dxr'*  i*i  the  ititi-r*  arlilaKinnii^  niu^<  1«  - 
the    (iiri-«ti.tn    tfli.    ninvf    tlic    sttimi'u    df    atta>     fmin    the    vi  rtf liral    colmnn    ah.    likr    Ih- 

txttrnal  tnti-rcnstals,  whuli  run  in  tin-  liirrffii.ii  kl.  During  thi>  .ImiMi-  aitifUl  tl.' 
r  nnist  lie  iip<  rt-asr.i.  Imaii^f  tin-  iiui-i  li  >*  I'f  tlu-  ni'i'pr  nitrn  "stal  s|iai  rs  »..ik  sinui! 
•  \    the    Mihr.'    th  irax    is    sliKhtIv    elcv.itcil    ilntniK    insitiralmn        FrcP.'    tin-    v.  h»-iin-    it    i- 

the  cxtirnal  inttTH'jilaK  nn.I  tlir  iiitmartilaH'iHMis  nn.isrlcs  mu^l  Uv  the  saiiu-.  (It-'I 
'Hill    /'/ivjri.>/,^.;v,> 


i 


The  Action  of  the  Diaphrag^m 

It  is  j)ossililc,  however,  that  the  main  function  of  liotli  the  iiitercosl;i' 
liuiscles  is  to  reu'ulate  the  tone  of  the  intercostal  spaces  and  so  pievcir 
their  suction  inwards  when  the  nejrativc  pressure  in  the  thorax  increasf 
'i.  e.,  suction  lieciniics  jrrcatcr  i.    The  ascent  of  the  lilis,  while  i)roducili!;  in 
increase  in  the  antcro])osteiior  and  tians\crsc  diameters  of  the  tliora\ 
Would  ilccri'ase  the  vertical  iliaiiicter  if  'his  was  not  counteracted  liy  tl  • 
lixatiiin  of  the  lower  lilis  ami  the  descent  of  the  diai)iir«irni.    The  jn-ripl 
eral  edu'cs  of  the  diaphra<;m  are  attached  liehiiid  to  the  luinliar  vertch'ii 
laterally  to  the  lower  cdtrcs  of  the  six  lower  ribs  ami  their  cartilafji- 
and  ill  front  to  the  tip  of  the  ensiform  cartilage    The  fillers  converjj'c  I 


Tin      MllllWns    n|-    l!l>l'll!  VllclV 


:!-M 


.lilrr  till'  I'cMliiil  tciiclciii.  ;iiii|  llii'  liitiTJil  bluets  ;iii'  pii'sscil  upwiifd  liy 
'Im'  iiilraiilHliiiiiiiiiil  |>ii>iti\i'  .iinl  iijtriitliiii'iii-i<'  iicjiitixc  picNsiiri's,  sn  tli.'il 
iliry  Inllii  ii  ilciiMi'-sli.'ipi'cl  \;iiilt.  with  tin'  lixcf  in  the  liu'llt  sicli"  aiiil  tiic 
-iniii.-ii-li  ami  the  N|)l('<'ri  in  tlic  li'I't. 

Kiii-iiiu:  cxpiiiiiinn  tlif  hiti'ial  imIui's  nf  tlif  iliapliiamn  arc   in   iMintact 

Mil  till'  parietal  piciiiii  nl'  the  tbniaric  i-a\it\,  Idriniiiy:  wliat  arc  kimwii 

'^  Idc  iili'uial  >iiiiiscs.     Diirih'j   iii>pii'atiiiii   tlic  lilicis  ot'  the  diaplirauMii 

■-liniicii;  this  Nti-aiuhtciis  iiiit    till'   ari'li    cil'  the   iliai>liiaL'iii    ami    pulls   tlic 

itciai  eilycs  (if  till'  (liapliiaijiii  ;\\\!\\    I'niiii  the  parietal  pleura,  thus  diicii- 

.1'.'  up  the  pleural  sinuses,   into   which   the   liinu's  dcsceml       I'siially   the 

•  licniiii.'  lip  1(1'  the  sinuses  is  acciuiipanicil   hy   a   slicht    retraetiipu   nl'  the 

'.lernal  clu'st   wall,  which   is  kmiwn  as   Lilten  "s  diaphramil   pheminiendii. 

Till'  descent   III'  the  diaphra'.'iii   may   pindiice  a   iiiip\  eiiiciit   iif  t'riun    Id  tn 

I  "i  iiiiii.  nil  each  side,  which  accniinls   fur  a   rallier  inipnrtant    Iractinii  uf 

lie   Miiuii f  air  eschaniic   liy    the   luiics.      The  central    portimi    nl'   the 

'liaphraiiiii    dues    imt    imi\c    iniich    in    imriiial    respiratimi.    Inn    in    I'lirccd 
-piiatiiiii   its  iiiii\  ciiieiil    ma\    he  cnnsideraMe. 

Because  111  its  allaehnicnts  tn  the  hiwcr  six  rihs,  th utractinu  ot'  the 

•li.iphraL'iii  tends  in  pull  the  iiijiririiis  ol'  the  rilis  towards  the  median  line. 
It   iimler  normal  cimdit  ions  this  mo\<'iiieiil    i>  iip|ioscd  hy  the  act  ion  of 

'i xtcrnal  inteicost.ils  in  iiiisiiin  the  rihs  and  e\paiiiliii<;  the  hori/oiit,il 

ii.iiiielers  of  the   ihoiax.   and   liy   the   Imver   \erleliral    muscles,   which    lis 
'ii'-  position  of  I  he  low  cr  rilis. 

11"     rililliri      jnnl     irhiili     II"     iliaiilirdilni     'iml     tin      i.rlinnil     inh  friishil 

'<'.'m7(x   lililif    ill    llii     iriih  iiiiii/    III    11"    liiin  r   jinii    nl'    II"     lh,,i;ir    (\    nt    sunii 

I"  I'lirlmii  I   jiiiiii   till    sill  ml  jiiiiiil   uf  iliin/iiiisii.      it    has  ccmr.-ilK    lieeii    held 

ii;it   the  coiiliaclion  of  the  iliaphr.ieiii  produces  ;i   w  ideninir  of  the  lower 

!'iil  ol   the  lliiiras.  Iieciuse  ill   its  ih  scent    it    pi  esses  upon  the  ,'ilidominal 

!M'ci;i    and    so    distends    the    alidoincn    and    pushes    out    the    lower    rilis 

1 'mI    this  iiiiLtlit    occur  seems   not    improliahlc   hut    lliiii\er-    has  fecciitl\ 

'""11  hy  eNpciimcntal  ami  clinical  oliscrx  alioiis  that    ;hc  llarinu   in   the 

■ -lal  iiiariiiiis  seen  in  normal   iiispir,-it imi  depends  on  olhcr  factors,      lie 

■     'Is  attenliiiii   to  the   f;ict    that    the  coiit  raci  imi   of  the   intercostals   raises 

' '■  rills  ami   iin-rc;iscs  tin-  anynlar  di\er!,'ence  of  the  siil stal    liordcis 

I  ills  w  iiji'iiiiiM-  ,,f  ill,.  ;ino|,.  iimde  !■>    ll isial  mar'.^iiis  ;it   the  tip  of  the 

■ilium    is    \cry    priiiiounced    in    paral.xsis    of    the    diaphra'jni    while    in 
'  ill N  sis  of  the  iiitcrcoslal  muscles,  the  costal  Inn  del  s  are  draw  II  tow  ards 
mi'liaii   line   and    the   siilieostal   aiojle   is   decreased,      'rjiis   sliows   tli;il 
'    'liaplir;iem  iiiiist   tend  111  diminish  the  aiiudc. 

riic  line  of  Iraetiiiu  of  the  di.-iphraem  is  a  sti-.iiirhl  oni'  Joiiiiinj  the  ecii- 
li  leiidnii  with  the  eilt'c  of  the  rilis  When  the  dia|ihr,-i;;iii  forms  a 
'  II  deliheil   arch,    it    e\eits   its   tr,-ielinii    at    a    disadv  anlau'e,    ami    tin'   e\ 


1 


i; 


t  I    ■ 


il 


Bit 

I 


»       !' 


I     -   I 


\i'2'l 


nil;  Ki.siiKATiDN 


I. 'ail    .1::.)     ..-A     1  ..-111' fi-     .■!     111.-      li.iiliraB" 


1  Nrrni.il    I'.'Ui'.ii    rit    ihaplKicii'        ^n-tal    matctiiH    niuVt-    ..tit    'ltiria«    iii-i.irati<'n. 

.'  Hik!i    pi.-iti..ii    .f   .i:ai  liiiiKiti.      N    *     -il    M,ii«.tiil    iii"vtui.rt!    ..!    ...-!al    !i!ar«in§  act.ni 

X  t.nw   i.'iHJIii.ir    nl    .ti.ii-lira«i!f       f'-^.i    nia^t't-i-    hi--\t    iit   -h-fhii    ii'-[-)r;iH'-Ti. 

4  \iry    t"W    I't.-ili'.ii    I't    -Il  i'''''arflr(        v  ■•*tal    iitarL'UT'    nuiv     "Mt    tlHrtitji    nupiratint: 

5.  Aaiial   liiif  of   tri. 'I"!!   (.1   ■haj.hr.uc"       (I'r.m   T.    \\ii'«ai.'    T-"''i  > 


nil;   Ml  (HANKS  UK  i:i>riiMTi(iN 


323 


I  'i.i^i.l::l    1-.    vlj..rt     t|i.     ,  h. 


^^lla.  inifnl.   .-i    ilu'   -,1  .[.Ir  ,kiii      •]    |l ,i.il 


■>.     .■  II    .l.innK    ii,<i.,i  .11..!, 
aimnuiit       ^'.-'trtl   iii.triftll   it"m  t-iiMhiriji    tn   niiiih 


N.T„„,|    ,...,,,..11    ,1    il,.,,,|i,;u-ii,       »  .i-t.il    ir,-,ri.M, 
l'"-iu..n  ,,1  ,l,ai.lit.i,;.ii   i,i  t,,,  ,.,1  ,.,r,!i.„    ,  ,.) 
V'>   tiiwanl   ntf,li.iti   hill 

i'-.Mli.m    ..t    ,),.i;.hr.,Kni     v     l,H-,lr.|    ,.,rl,a.     .  nUr^.  ii  .  lit        I  ,  f  ■     ,,,.|al    ,-v,t..„.    ,.    CxrH    "r 
'    i:i    •iuritid    iii'il.iratfiii. 
l'--lti„|l     ,.|-     ,li.n,,h,.-,KM      „,     ,,l:!i1   -1.1,    ;     .,,.1,.,,      ,1,1.,,,; t  1;..   I   1      ..,.  ,i     ■,■•,.,•■„       .     •   >,    1      ,r 

■   '"   'I'lriiiK  ,n'.|.ic,iliM,,. 
'  ■■Mil    n:,,,;,,,         I  I-,,.,,       ,■      \\  ,.  ^,,,     -I.,.,,,  , 


>'i 


^?^  vm 


fe*y-^ 


!»[RW^i^?«(?2»fK^ia<<£S''T«S; 


■.>''!??»  p'^^;-*; 


:11M 


Tin     III  ~l'll,'  MIDV 


Ii'in.il  iiitti cn^iii Is  |i;i\f  llic  iM;i>tci\  mill  ciiiiM'  tlic  cosliil  liiudiMs  |i' 
s|iii'.iil.  Wlii'ii  llic  ;iii-li  111'  till'  iliii|ilirii!.Mii  is  (li'pri'SM'd.  jis  in  iilcuri>i,\ 
with  ('lTii>-iiiii.  i'iii|>li,\  Miiiii.  mill  ciiiiivciiiii,  llic  liiic  lit'  tijirliiiii  aiul  tin- 
liiii'  111  till'  iiiiiM-iihir  liliris  111'  till'  i|i;ililiiiii,'iii  riiri-rspnliil  iimri'  clnsrlv  . 
Ml  ihiil  till'  iliiipliiiiuiii  is  iiMf  111  iisi'  its  full  I'lircc  ;i'_';iiiist   tin-  i'ili'ri-i>st;il 

iiiiisi'ji's.  willi  llif  ii'siill  thill  ll still  liiinli-r  mines  lnvvmils  tin'  mciliiin 

liiii'.  Till'  iiiivrs  nt  the  illiVi'iiiit  lilicis  111'  till'  iliiiiiliiiisiiii  \;iiy  irn'atl>  ; 
llii'  airli  is  iiiiii'li  li'ss  iiMikfil  in  llic  portion  atfaclicil  to  tlic  mstal  luar^'in 
near  the  iiiciliaii  line  lliaii  in  lliat  attaciitMl  in  the  a\illai>'  liiie.  l''o|-  this 
reason  the  aiilei  olatei  a!  part  nl'  the  diaphia'-'iii  reijiiires  less  ileprcssioii 
to  Lri^e  it  a  hiiri/iiiital  position  than  is  reipiireil  for  parts  oeciip\  iiii>'  .1 
more  lateral  position.  A  small  peiieanlial  elViision  or  an  increase  in  the 
si/e  of  the  heart  max  therefore  depress  the  iliapliray:m  siitlleieni  ly  to  irix  e 
it  liiaster\  over  the  inter. ostals  in  the  front  iiortieii.  so  that  the  costal 
liiiiilcr  iiia.v  here  move  towanls  the  iiiidliiie.  while  the  lower  Imrders 
miAe  ill  a  perfectly    imrimil   manner     see   l''i'.^s.   llli  and   117'. 

Diii-inu  fiii-iiil  liriiillniui  si-seral  nniscles  are  liroiiiiht  into  pla.\'.  ainoirj 
the  most  important  of  which  are  the  sealeni.  stcrnoiiiastoid.  1rape/iiis. 
pectorals,  rliomlioids,  and  serratiis  matrniis 

'I'll  ere  has  lieen  ciiiisiileralile  deliale  as  to  w  hither  1  1  /iif'il  lun  is  iiormall  \ 
an  active  or  a  passive  pro. -ess  I  ■|ii|oiilitei|lv  the  expiratnrv  |ihase  iiiider 
normal  conditioiis  does  not  leipiirc  the  same  miiseiilar  clTorl  as  docs  that 
of  iiisp:;atioii.  lint  there  are  manv  ohscrvatioiis  which  indicate  that  e\ 
liiratioii  is  p.irtiv  under  miisinlar  i-ontr.i!.  The  ididoniinal  niiiscMlatiire 
iore\ample,  increases  in  tmic  diirini.'  expiration,  so  as  to  lirinir  alioiit  a 
rise  ill  the  ;i  III  luminal  pi.-ssi|ie,  with  the  result  that  t  lie  iela\eil  diaplira'jm 
is  piishcl  lip  into  the  thoracic  eavitv.  'Co  this  e\teiil  at  Nast.  expiration 
is  a uiipaiiieil  \>\     iicreased  miiscnlar  adivitv. 

I'.el'orc  h'av  iiiir  the  siilijcci  of  the  diaphraiiiiialic  movements,  referenc' 
must  III-  made  to  the  recent  ohservatiiills  of  I.ee,  (I'uciither  and  Melelicv 
lieariiiv'  oil  the  irciieral  pliv  siolou'ic  pinpeilies  of  the  .|iaii!ira'_'iiial  ic 
muscle.  They  point  out  that  most  skeletal  iiiiisi-les  in  the  iiviiii;  liodv 
.■oiitraet  with  varvinu  dei;rces  ..f  Intciisiiv  and  .it  irregular  intervals. 
Iiclvveeii  which  relalivelv  loii'^  periods  of  r.'si  oc.-iir,  Init  the  diaplirau:iii 
Iiiiiii  liirth  to  death  performs  ;i  coiit  iiiiioiis  succession  of  hrief  contrae 
lioiis  of  fiiirli  re._'iilar  rhvlhni  and  uniform  extent,  alternatini;  with  lirie! 
periods  of  rest.  lis  iniiselc  tilicrs,  together  mIiIi  ihosr  of  the  other 
respiratory  muscles,  therefore  hold  a  iiniipie  position  aiiionu'  skeletal 
muscles,  which  sui.'._'csls  ii  .rude  anahnjy  with  that  of  the  he, lit.  Tin" 
have  coiiiparcd  the  pliv  sioloL'ic  properties  of  the  diaphiii.jm  with  tho^' 
of  the  extiiisiM   lon'.jiis  ili.jitoiiiiii.  the  sartoriiis.  and  the  sulcus,  ami  I'ouii'i 


Till.    MlrnWHS   OK    HiisriHAlKiN 


:{2:> 


iit  till'  iliii|ilinit.'iii  is  n|iiiM'il  III'  a  mui'li  inorr  t'tliciciit   iinisciiliii'  tissiir 


tliiiii  tliiil  III'  tilt'  otlicr  mil 


The  Effects  of  the  Respiratory  Movements  on  the  Lungs.    Tl I 

|iiui|iiiT(l  ill  till'  iliiiiin>icp|iN  (if  llic  liiiiLTs  liy  tlic  iiis|)iriitiii'y  cxpiiiisioii  nl' 


laiiTi- 


III'    Tllnlil 


•ii-  i-avitv   ail'  liiil   iiiiifiinii.  sin litTcii'iit  ))arts  of  flicsc  striii- 

Hues   an-    nut    ciiuallx    rxti'iisililr       l'''iini    an    aiiatHiiiii-    >laiiili«i)int.    tlic 


MIL'S  iiiii.v  ill'  i\\\  mIi'iI  into  tliii 


I    Tl 


11'  iniHT  (ir  iiint  ziijic  I'lmtiiin- 


in    till'    liriiiH'liiis,    art 


rr\    ami    xciii,    am 


I    tU' 


ir    main    siiImIiv  isioiin 


Tl 


ir 


'.ii-l:i'  aiiiiiiiiit    111'   lilnniis    lissiii'   in    tliis    ic.'iuii    uIViTs   liirat    ii'sistaiii'c   In 
ii>  i-xiMiHiiii-i  I'lii'cc.       'J     '['lie  intcrmciliati'  /itui\  riiiitainiii!,'  tin'  vasi-nlar 

iml   liiuni'liial    lamilirat  mils  radiatiiiL'  Inwaiils  tlii'  siiil'ai I'  tlic  lllM<rs, 

"itii    pulliiKliai'.v    tissue    ini|ilalit<'il    lii'tuci'ii    1  lie    la.vs       Tliis    part    nf   flit' 
iniiu's  has  \aiviiiu'  ilt'u'ri'cs  nf  I'NlciisiliilJty.  tin'  |)iilmnnar.\-  lissiii'  lia\  in:.' 

!liv  iiiiist  ami  the  \asi-iilai-  ami   Ini Iiial  llii'  least         :'.     Tlic  mitcr  zniio. 

iiiiliaps   '2't    111    :!()    mill,    in    ili'ptli.    i'iim|iiisi'il    .if   pnlmmiaiv    tissue    and 
"piallv    eNtensilile   Ilii-iiiii;liiint      Keitli'  'I'lii     e\paiisiiiii    uf   llie   liiim   is 

.iiTiilliplislieil    li\    a    lilnviliu    apart    nf   llie    less   exteiisihle    ra  \  s   nf   tissiii'   sn 

■1^  '"   pi'i'init    tl Npaiisiiin   nl'  the   nmre  esteiisilile   ))iiliiiniiar\    tissue   lie- 

U\ici'   iheiii.      Keith  eiiiiipales  the  iiieeliaiiism  tn  that   seen   in   the  iipeniinr 
'■I  .1  .lap.inese  fan, 

lii'i-aiise   the    liiii!.'   expamls   in   the  ijiiei'tinii    nf  h'ast    resistani'c,   stmlv 
•I   till'  inllateil  ijeail   liinu  ilnes  imt    rcMal   tin liial   eNpaiisimi   hrnnirlit 

.ihnllt    In     the    ihnraeie    llim  elllellt  s.        In    the    lixilie-    linilv     eXpallsillll    is    mule 

iiiiiiteij  in  siiiiie  ri'iiiiiiis  than  in  ntln'is.     <>\   the  li\e  areas  >v|iieli  may  he 

'listiiiu'uislieil  nil  the  siirfa if  the  liiiijis,  three  are  In  i tact   witli  rela 

■|\rly    iiiimii\alile    parts   nf   the    ehest    vvall.    ami    therefnie    eal t    he    i'\- 

I'aiiijeil  liireetlx.  These  ale;  the  iiii  il  iiisl  i  nil! .  ill  I'niitael  with  the  perieai' 
'Inilii  a  III  I  the  si  I'l  let  I  ires  111  the  Ineijiast  iniim  ;  tlli'  il<,i->,il  vk /■/.(.  <  .  in  eniitael 
Mill  the  spiiial  rnliimn  ami  the  pnsterim-  aspeet  i<\'  the  tlmraeie  ea^'e,  ami 
'lie  11)111-, il  Mirliiii  .  The  miiliniis  nf  tile  first  pair  <<(  rilis  ami  the  maim 
'"'Uwn    e\paiiil    ehielly    the    allterini     at     i     \  elil  rnla  tela  1     part    nl'    the    ape\ 

I  the  liiiiLT,  aiiil  lia\ ly  an  imlire't  inlliienee  mi  the  ijnrsal  part  nf  the 

l"\      i   <•..  the   part    lyiiiL'  ilireetlv   in   Innit   nf  the  iieeks  nf  the   liist   ami 

■I'nml    rihs,   the    must    enliuiinn    site   of   plllmnliary    t  lllieri'lllnsis.      The    twii 

-irlai'is  ,i(  the  lniii;s  whieh  are  ilireetl\    e\paiii|eil  are  the  liiaphrau'liiat  ie 

Mil  the  \  eiiti'iilateral   nr  si,  rnnenstal,      .Melt/el'    fiiiiml   that    the   iie'.'ative 

iiie^sin-e  ill   the  tlinra\  ihirini!   iiispiratinn   was  least   ahni^   the   relatixi'l-. 

^''' :ii>    "alls  nf   the   thnra\.   ami    '.nealesl    In    the    reeiniis   nearest    the 

'  ■I'lii'i'icm.      I'riiiii   this  he  i-niielmles  that    sniiie  i>\'  the   e.\pamlinir   foree 
•  i"sl  as  it  passes  thininj-h  the  lini'^s  tn  the  siiffaees  ,\\'  imlifeei  expansion 

M:ih\     nliservers    |ia\e    elailiieil    thai    the    expailsinii    nf    the    |iiiim    ,hies    mil 
'■"  '■   jilaee   thrniiul t    iiisi  a  ii  I  a  iieniisl  \    ami   e.pia!l>        This    j-,    il  hist  rateil 


i   !l 


:!•_'(; 


Till:    Hl>riKATl(i\ 


t  '     :! 


I 


•1: 
1   iJ 


ii 


If 


till'  I'iK'l  th.'il.  ill  tl 


i'('i:i<>M  iiiiiiK'iliiitrlv  >iii'i'<iuii<liii^  a  lix-iili/cil  n>ii 


Miliiliiticiii.  ;i  lliiiil  lias  iiHMi'asril  icMHiaiii'i',  wliicli  XMHild  imt  lie  the  ca^' 
il'  till-  iilaxaliiiii  pi'imIiicimI  was  ('<|iially  ilistriliiitcd  tlinitiulnmt  the  luii'j 
Tlie  riiiit  ol'  ilic  limtr.  wliii-li  lias  tri'iiorally  bccii  reifardi-il  as  iiioro  m 
less  (ixrd.  iiiiili'ifiiics  in  iiiiinial  linalliin^t  a  definite  forward,  dowiiwaid 
and  i>iit\\aid  iiiii\ cini'iit.  and  llie  heart  shares  in  tills  iiioveiiieiit  (KeitliK 
The  tiMiN  eiiieiits  ni'  the  lower  lilis  and  dinphra'.nn  are  ri'spoiisilile  for  tlii 
e\|iaiision  of  the  lower  lilies  and  dorsal  portion  of  the  upper  lobes  ot'  tin 
Iniii."".  whereas  the  iiio\  i-nient  of  the  upper  live  rihs  expands  the  aiiterioi 
portion  of  the  upper  l.ilies.  The  relative  iiifreipieiie\  of  pleiirltie  frie 
t ion-sounds  and  pain  over  the  upper  lohes  as  compared  with  their  fie 
ipieiiey  o\  er  the  lower  lolies  is  e\|>lained  liy  the  fact  that  the  expausioi; 
of  the  upper  lohes  is  aeeomplished  with  little  di->plaeeiiieiit  of  the  pleura' 
siirt'aees.  \\  liereas  in  the  lower  lohes  cxpatision  is  aeeompanied  liy  a  \s\\<\ 
iiitr  of  the  lnn''s  aeioss  the  rihs. 


i! 


CIIAPTKH  XXXVII 

TIIK  CONTROL  OF  TIIK  KKSI'IRATIOX 

The  piiiticipfilioii  of  Midi  \vi(l.>>,|)rr;i(l  moiips  of  inuscles  in  the  respira- 
iniy  act  (loniuiids  that  soiiu'  nicchaiiisiii  lie  provided  to  insure  its  adef|iiate 
.  uiitrol.  With  every  inspiration,  for  example,  the  museles  of  the  nln- 
ii.isi  act  so  as  tn  cause  liilatatinii  of  tlie  iiai'cs.  tlii'  vocal  cords  are  iilt- 
(I'lcted,  and  the  intercostal  niiiseles,  aloni;  with  the  sealenes  and  the 
■  liaplirau'rii  ate  cuiitractiiii.'  \\hilc  the  muscles  of  the  ahdomiiia!  wall  are 
rclaxinfi:  and  all  tliese  e\ents  occur  at  exactly  the  proi)er  time  so  as  to 
'liiiir  ahouf  the  most  eftlcient  oi)cnin<r  up  of  the  ihoiacie  <'avity.  Kvi- 
'li'iitly  there  must  lie  s<iiiic  mechanism  to  insure  this  perfect  control.  This 
I-:  cfTected  throuirh  the  nervous  system. 

THE  RESPIRATORY  NERVE  CENTERS 

The  efferent  fihers  to  the  \aiioiis  L'loups  of  muscle  originate  in  their 
"^pecti\e  motor  neurons,  «hich  in  most  eases  are  situated  in  the  pray 
iiiitlcr  of  the  spinal  cni(l.  The  liariiionioiis  action  of  these  motor  neu- 
"iN.  or  suhsidiary  center-^,  is  hrou'rht  ahout  liy  the  transmission  to  them 
i'  impulses  frojii  a  hiiihcr  or  master  center  jtlaecd  in  the  medulla  oli- 
'iii.'ata,  the  i);ith\va,\  of  transmission  lietwcen  this  master  center  and  the 
ilisjdiary  centers  lieiiii;  in  the  lateral  coliinins  of  the  spinal  eord. 

Tiie  evidence  that   fht   chiif  r< ^jtirntorii  niitrr  is  in  the  medulla  is  fur 

-lied  hy  oliserviiiu'  tlie  cfl'ects  ))riidiiced  mi  the  respiratory  movements 

'■    serial  destruction   of  the   cerilirospinal    axis   from   ahove   downwanl 

!'     this  method  the  approximate  position  of  tlie  center  is  found,  its  exni't 

'•ation  heinir  then  determined  1>.\   punctit'oiiii  destruction  or  stimulation 

'   tlie  supposed  locus  of  the  centi'r.      If  Me  destrov  the  cerehriim  from 

I'li-e  hackward,  piece  hy  ])iece,  we  shall  liml  that   loi  marked  effect  is 

■'Hliii'iMi  on  tlie  respiratiuiis  until   we  arrive  at   aliout   the  middle  of  the 

■iliilhi.  when  immediate  |iaral\sis  of  the  respiratory  movements  occurs 

!■    \.  e  now  priiceed  to  puncture  \ariniis  areas  on  the  floor  of  the  fourth 

nlriele  in  another  animal,  we  s|i,-ill  lind  an  area  calleii  tlic  iiiinid  rital, 
"■•iteil  aliout  the  tip  of  the  calamus  scriptoriiis,  destruction  of  wliieli 
•lUses  immediate  cessation  of  respiration  It  is  lieliexfd  tliat  the  center 
■-i'lis  in  the  <rroiip  of  ner\  > lis  kno\\  n  to  neiiroloirists  jis  the  fasciculus 

iit;n-iiis.      Tt    is   liilateral. 

■.\-27 


'■  ;■! 


1 
it 


f  1 

I! 


•MM''>^ 


il  11 


\2!< 


I  III    III  -nil  M  iii\ 


•//..   >,//.>„/ 


iiir'i  I  I  nil 


I  >■  iilc  I'liliiclv  ilr|(i'llilc1il  ll|>ii|i  llii'  iiiiislci  cclil. 
I'm  llicii  hilt  liiHiiliiiis  iiclioii,  lis  IN  nIhumi  Ii\  (lir  liifl  thai  if  the  phlilii 
iiicitni'  iicMiiiiii      whirh   is  siliiiiti'il  ill  thf  ii'i\  ii-iil  spinal  cnid  hrlwivii  11 

fciiiith    an. I    si\th    spinal    sfjiiirnis      is    isnl.it.'d    I'l ih,.    iiM'ilnlla    l..\ 

lali'ial    hiiiiisi'i'iniii    nl'  thi d   jiist    ahnvr    iIh-    Iniiilh   sciriiiciit    ami    h 

iiM'sial  sfi-tion  nl'  tin-  ■•mil  uppnsiic  thr  rcnli-r.  llir  i-ipi-icsp.mijiiin  iialt'  ■ 

the  iliaphiaiiMi  1111  |.iii!.'iM-  pall  icipalcs  in  the  inspii  atui  \   art   i  sec  |-'ii;    ||s 

'I'lic  ihiiC  ci  iilii    on  cither  siijc  nl'  the  iniijliiic  nl'  thr   nirdiilla   is  cm 

l''i|    «illi    llii>    HM.Inr    in'Uiiins   oi'    l>i,tl,    si, Irs   nl'    ili,'    spih.il    lunl.    .is    ; 

piMM'cl  li\    ihi'  rolluwiirj  csiKi  iiiii'iil       Whrii  the  c-cntial  cinl  nf  the  \ai:ii 

iii'iM'  is  si  iiiiiilali''!.  till'  I  espiiatiiiv    i-cnti'i   I nun's  cM-itc'il  ainl  the  ii'sp 

laliiiiis  nmii'  piniiniiiii-cil,  the  pail  ii-ipal  inn  nl'  the  iiiiiscirs  mi   Imih  si,|. 
nl   the  limlv    licini;  rcpial  ill  i'\lc  lit        IT  11  nw    u  !■  I  Msec  I   I  In'  inn  I II I  la   (hn\  n   1  i 


» 

;■ 

.  ;  / 

H     I 


1  . 


I IJ    11-     I' 


.1  l...    1>  .'.ll     •      t   li 


111.      .  -Mlt 


Ms    y 


mil  Ili  lie  ami   lepra  I   t  In-  ^I  i  m  ll  l,il  i. I   niii-  \a'_'l|s,  ihi'  iiiiiseles  mi   hnth  siii. 

will  still  pai-tieipale  III  ihe  III.  leave, I  r.-pirat inn.  wln.-li  tli,'\   \\t\\  likex\i>,- 
il'th,'  e.T\i,-al  e,,r,|    is  l,is,.,.|,.,|  ,„    ||,.|iiis,M-t,',|    Iml    ih,'   iiii',|iilla    li'l'l    inl.e 
I  I'i^e    !  I'l         The   siiii|i|,-si    intri  |n-,'lati,i|i   ,,1'  lli,'se   r, •suits   is  thai    i-,,|iini  ■■ 
siiial   liliei-s  eniineel    Ih.iIi   haK,s  ,,|'  th,'   ii  spii  atniv    ,-i-iiter  in   the  Mieijn! 

ami  that  ,m,-Ii  hall'  is  ai-,  ,-n •■•ie,l  \xilh  the  iimlnr  iienrmis  nt'  l.ntli  se!' 

nl'  till'  spinal  i-ni-,l.     iM'li'ii,  es|ie,.iall>    in  \,iuii,j-  aiiiiiials,  a  lieinisi','ti,it;   ■ 

'I nnl  ,'aiis,>s  eessalinii  nl'  tin-  liin\  ,MiH'iit s  nl'  till'  ili.i phrayiii  on  the  sa'i 

si, I,':   liiit    this  paialy/e,|   si,|e  at   ,.ii,-,'  lieuiiis  tn  I'ontiaet    aL'ain    \\  hen   ' 
phieiii,-    nl'    ill,'    oppnsii,.    si, It     is    ,-iit.    pi-,.lialily    li,'eailse    tli,'    resiiira' 
impulse  ili's,-,'n(lin.j    trnm   th,'  ,-liii-r  I'eiiter.  on   limliny;  its   wiiy   ;ilniiy    ■ 

lor  I'eliti'r  o|'  tin-  sam,'  si,|,-  ,,1'  tj i,}   lihieUe,!,   is   l'ori',',|   t,i   l'onn\>    II 

ern--s,-,|   path.     'I'll,'  i-rnssiiiLr  in  th inl   is  lielii'\,'i|  to  take  pl.ee  at    I' 

sail..'    Ii'\,'l    as    ihal    al     w  hi,-li    lli,'    siilisi,|ia  i  \     e.-nt,'!'    is    Ini'aliil    .  W 

I'nill'l 


li!  1 


I  III     I  IIS  lliiil.    liK     I  Ml      111  >lll;  M|ii\ 


;li» 


i|||i'nIiiiii   iinw   iiriM-  ,1,  t(.  Ii„,r   lln    rin,  I    niihr   i iniiliiiHiili  <       !>   il 


|iilirlv  li'llrN   ill  ill.' 
I  i,iii>liiis>iiili  111  jl  11* 

III    till'    SCllM'    llijil     I' 

lMlinitirit\     III    llir    fi 


l.il   It  i|i|niii|s  I'ni-  j|<,  itctivily  I'lilircly  mi  tin 

Us    illl|'ll|si  >    ||M|ll    1  Im'W  hrl'.',    Ill     is    it    initillMMlii 


III, 


UK    lllllrpi'l 


lll'llt  l\      nl     >Ul|l     ihlplllM'N  .'        Til 


I'll   I    .I'liki's   it    SI  iiii    imi    iiii|ii'iili;ilili'   th.il    the   cciitfr 

lii'll    ciilitliils    till'    , nliliMti'    iirtiiili    'il    till'    li'N|iii;iliiry    lilllscji's    wiiulil 


,U.i  I 


iImi  liiiv  I'  nil  mini 


lilt    nr  iilltiilii.il  |i'   I'liH  ti' 


'I'lir  .i'-li\  it  \    111'  --lli.l 


I    .ill    until' 


liiitii'    lf-|iiliitii|  \    iM  lilii     viniilij,    u| 


iiiiM-.  It  >iili.|ii-|    til  i.'iiiil    \  iiriiitimi 


.1    ll'sllll    111'  i-|llHI!.'is    ill    till     i'<i|il|iiisit  j.iii    dl'    III,'    IiIiiimI    sl||i|i|y  iiijr    jl     ,11,, I 


iIk'    I'ili't    llinl    il    vt  lis   lllltnlll.it  ic    w  iilllil 

il  I  \  mis  iiii|iiilM  ~       jiiilii  i|   il 


liiil    !!■ \f   it    rinlli   till'  illlllliiirc   III' 

^il'li  111  i'.i||rri\  r  iif  l(ic  .i  l|  t  il|||ll  I  il'll  \ 
"I  till'  liiltii  ;|s  liiiliM  ,,)  ;i  In',  ii|,|,'|',  1!  illi  Its  imiiiiil  IliliitiiillJllfi 
ll'iil'lrllt     ll(ln|,    allVnill     Ml  Ml     illl|il||sis         lis    ili|t..||i;itirit\     llii^llt,    tllCII. 


\:<       11'        III, 


■'      "       "   "■""      I"    r-      IH      111 

ilii"'il    ;-U    .li.liiilM'itf    III. 


1,11..    .ui'l     ;i|'|., 


iM-    I.  -i.,r,ili,. 


"I    llii.    -I  I.     imII 


't     nil.  II.  I.     «illi    III. 
1...11I1M11;     mi     l„ 


I  ti,.ns    i,,,ii,     ^t  j.,i) ., 


i.lv    »ill 


■I    \.\.   1-    -II"  'il,il,"i        II     I'.'lli    -..  II. 
I'll.  1 1.  I.  >l     uilh     Mil     !!i.      -I't.      "I     ih. 


iMiii'lv    il    r;ii-|iii'    ,,f  siiTi'tv     i-iilli',|    iiil 


1   |iiii\    iiiilv    w  Inn    til,'    iiiiliii'iiii" 


i-r  ^\  I'l  I-   Im-  SI. Ill,'   I  ciisiui   I  I'll 


.riiiiiiil\    .  iiiiliullini;   till'  I'i'iil 

till'    l|llrsti,i||     wliicll     ,'it      |l|','s,'llt     ciilltl'lillls     Us.     h 

Mil'!'  iii.'iy  111'  iiiiiv    in.t   iii'l   iiiitiiiiiiitiii-ilh  ,      M 


lUI'M'l',     Is     W  llctlllT     il 


itiiUfii   t,,  test    this 


Is'ililtlllll     111'    the     ITllll'l' 


il-     Till 
III 


lliv   rS|ii'l'lliiflits   liil\  |.   1 II 

II    111    tlirlll    ilr|ii'in|ilii;    i||iiiii 
l|ilrti'ly    ;is    |i,issili|i'    finin    iitVrli'llI    lii.|'\i' 


11 t,   the   llillllli'   iif 


11'    llliist    si|i-i-('ssri||    i'\|ii'iiiiii'iit    liils    lii'i'ii    |i,'t|iii'iiii'il    ;is    lull 


iiilliii'iiiT  111    till'   lll'jlii'l-   lii'i'M'  i-,.|iti'l's   Wiis   I'l'liniMil   li\    i-uttiliu-  ill'lil 
jii  illllirli's  ,,f  th,-  <'i'r,'l,i'iii,i   ,,i    ill,.   |ii,|is,      'I'hi.   Ililllii'lii'i'   iif  iilVi'Iclit    it 


tl'ilM'lill.J    U|l    111!'    spillill    •,{     i\;|s    II'IMUM',1     |,\     (',  i||||ll  I '  1 1' I 


\     Sl'\i'l|l|.. 


-|i|li;il   nil 
h.'sti'i  i.ii' 


ij    lii'liiu    thi'   li'M'l    III'  llic    I 


llll  rlili'    IliM  \  .'s    ;| 


llll      s,',.(ii,||i||ir     ;i|| 


II'    Sli|S.ir\      Slllllill      I, lilts     111      t 


n I",  Hill    mill    iiliiiv  ,.   1 1||.   1,.\  I 


i-ii.iii      TI 


iKn  i-iii  t 


\V    V.iL;l    Hi'lc    ,'l|sii    ,-||l    In    |'i'|t|ii\i'    till'    Illl|lll 


ls,.s    1, 


n 

FFf 

! 

k 

1 

1 

B 

3' 

1 

s 

ii 

1 ' 

^♦i 


;t:iti 


illi:  Ki  -III;  \rii'\ 


i\   tliriii  til  till'  ii's|iiriilii' \  ri'iiti  I-.      |l\   Micli  an  ii|M'iatii'ii  lln>  only  lov 


I('s|li|';i|.i|  \    hi'lllnns  li'l'l    illlllrl       II' 


il'  till-  pliii'iiif  iii'rvf,  NO  that  tl 


ii'»|iiialiiiy   iiiiiMiiiciits  lliat    ,11    n.    ;ii,.  |iii>..ili|i'   aiT   lliosc   in    whii-li   1I1 


■  liaiil 


iia'jiH  iiarlicii>ali 


il  tlir    iiii'«i'lfs  ill'  tli(>  alu'  iia>i  ami  larynx.     \' 


\\:\-<  rnuhi!  lliat  tin-  aiiini.il  at't'M    il |ii'i  .itinri  wi'nt  mi  rt's|iii-in<r.  tlidinjl 

iiii|Hrl'ri'tl,\ ,  ami   that    tliv   1  ispiiatiMns  Minn    lu'cainr   itmi'i'   iiiarkctl   aii' 
aspliv  sial    ill    cliaiafli'i .     imlii'ii' i'li:    that    tlir    lilimil    was    nut    licrniiiin 


W'Pj 


'7i''i\ 


S  Pnrenicul  < 


Duphr 


li,-      1-'         PiaKiu-i 


-I.'->i     ■.\!i<i«     .   ,1-    ;n«     !i,.i  !t     I.)    ixi.:.ii.     111.-    tlinf    ii -i.ir.iloi  >■    « '  nl«  I     I 
afTf<  rtrit   imp^iKf*. 


|ii.,|Mi|\     iM'ialiii    ami    liial    ihr    i-hriiiHMi    rhaii^ri's   nci'iui'ilit;    'H    '•    ^^ '     ' 
ai'ti'"j  ilii<i'tl\    ■•11    ihr   !•( 'I'rr.   ^t  iiimlatiiiir   il    tu   L:iialt'r  ai'tivity.     '11 
I'liiirliisiiiii    sr.u:^    w  ,-ii  rail'' 'i   that    thi'   i'i-.|iiraW'ry    i-fiitiT   i-aii   act   ant'' 
iiialic-all.\  .  I'nr  the  •>iil>    |i'.-^ilih'  alTi'i'.'iil   iii'im's  h'l'l  in  tlic  ahnvo  prcpaia 

tinii  wiTi'  thiiM'  raiiird  tii  thi iili'i'  liy  thi'  til'th  mitvi'  (  l-'iLT.  I2I>). 

That    thf  I'l'spii  atiii\    c-mlir  i-<  cxtranidiiiatily  srnsiti\e  to  chaliKOs 
•hi'  (■(iiii|insiliiin  lit'  llir  hliMxl  lluwint.'  tlMini'.'h   it    i>i  a    t'ai-t   that  has  hoi" 
kiMi\\ii      '!■  a  loll'.;   liiiH-.  hut    il   i^  iiiilv    \'.  ithin   m'i'iit      rars  thai    tlu^  cxai' 


nil    icisKiui,  UK  nil,  UK>ni(\iiiPS  :{;il 

nature  "I'  lliis  ciiiitnil  iiiitl  till'  ii'Miiirkalilc  H«'ii.siti\  \U  i>(  tin"  i-ciitcr  t'lwiuils 
ii  li:i\r  hcfii  tl  'loiiuhly  csliililisliid      W(>  hIkiU  ntiiiii  to  fliix  iinpiiriHiit 
-ulijii't    Inter.     Mi'iiinvliilt'  v  •■  nIuiII    prncffd   to  cxiiiiiiiii'  tlir   iiiiiiiiifi'   in 
liii'li  the  ciMitf"  is  a(Ti'i-tc(l  li\   M'liNoiy  iiiipiiNt's  traiisiiiittcMl  to  it. 


3 

-3 


f 

3 


THE  REFLEX  CONTROL  OF  THE  RESPIRATORY  CENTER 

Till'  iilTciriit  iiciM'  liln'is  '.'oil!','  to  ilw  rrHpiratory  ci'iitcr  Tiiny  coiivcn 
iilly  III-  i|i\  i.licl  into  tw  o  mi.iips:  tli"  .•  w  liii-li  act  on  it  only  occasioiiall>  , 
'III   lliiiM'   \\  liii'li  art   nil   it    liioic  or  li's;.  I'olitilllloilHh'. 


The  Occasionally  Acting  Impulses 

To  till-  lirst  j,'iiiii|i  lii'liiiiir  alViTiiit  iicim's  iiom  piarlii-ally  cMi'y  i)Mit 


I  PI  II IV. 


That 


iiiipir^^iipiis  t'riiiii  tlic  >.kiii  alVi'i'f  tlic  ifspiiatory  cint 


;^  \M'l!  Uiiow  11  liy  till'  iiiiMiMMil  lni'atliintr  caiiMil  liy  ap|ilii-atiiiiis  of  colil 
.iti'i-.     Till'  iiidiiriii-i'  of  tln'sf  alTfi'iiii   iiiipiiKi's  is  often  vci'y  iiinrki'il. 
Mil  is  l'ii'i|iii'Mtl.\'  takrii  ai|\  aii'a;.'!'  ol'  in  stiiiiiilalinir  a  nculMirii  infant  t" 
Ivi'  till'  first  lircatli      Stiniiilaiion  of  llic  ti'i'ininations  of  tin-  fifth  ncrv 
till'  iiiiicoiis  iiiciiiliiatK'  of  till'   nose,  as  liy   iiihaliiii;  a   piini;oiit  mlor. 
'iMiiiiliati'ly    inliiliits   respiration.     To  tliesi'  oeeasioiially   aetinjj  affereiil 
!|iillsi's  may  lie  ailileil  the  iiiiplllses  that  are  conveyed  to  tlic  rt'spirator.v 
I  liter  from  the  hiirliei'  ner\c  eeiileis  of  the  cerelinmi.     These  imimlses 
I  e  laiL'clx   \iiiiintary  in  nature,  ami  enahic  ns  to  hohj  oiir  lireath  at  will. 
"^   III''  of  the  eereliral  inipiilses  are  however  also  involuntary,  their  exist 
me  lieiii!,'  seen  li,\    oliservin!.'  the  respirations  of  an  animal  hefore  ami 
Mel'  sei'tioiiiii!.'  the  polls  or  peilnni'le^    The  respirations  for  a  lime  at  least 
■i-niiii'  ilistinetly  alTecteil,  Init  lliey  later  retiirn  v^illi  perfect  re<rulai'it.\ 
I ';•  \  iiia.\  lieciiiiie  \<]y  ii  rei,Milai-,  hii',\c\er.  if  the  \a;.'i  as  well  as  tlie  pons 
'■  cut.     Other  experimental  exiileiice  of  the  existence  of  eereliral  respir 
.i'"i-.\    liheis  is   fiiriiishcil   li\    cei'eliral    locili/ation   experiments,     niirinti 
■iiinlatioii  of  the  eereliral  cortex,   for  c\,iiiiple,  a  marked  effect   on  the 
-piratory  iiiovcments  is  often  noted. 

l.''>piiatory  ilivthm,  unlike  that  of  the  lieait.  has  often  to  he  mnditieij 
■■i'i''r  that  the  respiratorv  mechanisni  may  lie  used  for  other  purposes 
III  the  \entilatioi.  of  the  liiii'js  This  jilteratioii  in  rli>thm  may  tak'' 
'If  1111111  of  a  mere  inliiliition,  such  as  the  ad  of  sw  alln'^  ini.' ;  or  tin 
-piiation  may  he  altered,  as  in  phoiiation  and  siiiu'iiifj.  .More  coiisiil 
ilile  alteration    in    the  expirator.v   discharsre   occurs   in    couudiini:   ami 

■ /iiiir,  and  still  moie  in  the  acts  of  mietiiiition,  defecation,  and  partiiri 

'  "I!  We  must  eoiiclude  therefore  that  the  rhythmie  stitniili  sent  out 
"III  the  respiratory  center  are  so  w.'ak  that  stimuli  from  other  souices 
■'}   iiislaiitly  iiiliiliil  or  clianire  their  form  at  aii\   stau'c  of  the  c\cle. 


ill 


H 


I  .*  i| 


MICDOCOW    RESOLUTION    TEST   CHART 

'ANSI  ond  liO  TEST  CHART  No    2 


1.0 


1.25 


2.2 


.8 


'•4     IIIIII.6 


j^     APPLIED  INA/1DE 


2SS      'I'iH'J    -  t\]. 


Till:    UIM'IUAIIUN 


!l 


iM: 


ill 


iii:!! 


i  i 


\-  •.', 


•'^•'       lii'i I  llii'  I'liiliiiiis  111'  ihc  yliiNsopiiiirx  iiyi'iil  nci\c  inhibits  it- 

I'ir:  I.  wliicli  cxiiliiiii^  llic  ImMiiii;  <,]'  liic  lu'ciith  tluit  ucciirs  in  sujii 
lii«  iii'j. 

The  Continuously  Acting  Afferent  Impulses 

''' ntiiiiiiiiisly  jicliiiii  iillViriil   iiiipulM's  swr  liiinsniitlcd  to  tlic  cliir: 

ic>|iiiiilci\   <-ciilfr  liv  III)'  \;iui  Jiiiil  their  hi-; hcs.  lh(    superior  hn-yiiu'eji : 

'ii'i'\es.  \\  ihe  \;ii;u.s  iier\e>  ;ire  eiit  iir  their  e(iiitim;it.\  severed  h\ 
li<'(/iii?  ;i  pdrtinii  (,r  ihei'i.  the  respirjitcirv  muxemeiits  iiecimie  iii!iriieill> 
^hi\ver.  I';\  ich'litly.  Ihe  \;iyiis  nerves  in  sciiiie  way  iiilirv  up  tlie  respir:i 
l'ii\     iiMi\eiiieiits.      Au;iiii.    if   tiie    eentriii    end    iif   either   Villus    is   stiiiiii 

'•■''''1    "i'h   tl idiiiiiry    inteiriiptcil    I'liriidie   eiirreiit.   a   proround   elTiTi 

on   tlie   i-espiiiitoiy    iiupn  eiiieiits   is   iisiimIIv   iihseived.      This  elTect    is   liuw 
ever  not   strictly   piedietiilih'.     I'siially   there  is  a  ipiiekeiiiiiir  of  res]iirii- 
tiipii,  anil  if  th.'  stiiiiuhis  is  a  stnuiM-  ,,iie,  Mieiv  may  he  a  standstill  of  tie 

•'""'•'i'^    ill   the    inspiratnry    positi'iii.      On    the   cither   liand.    if   tin iitral 

end  cif  the  iierxc  is  st iiiiiilatiMl   with   other  types  of  stiiiiiili.  as   hy   slow. 

weak    laradie  shocks  or  h,\    the  stiimilus  produ 1   liy   the  closure  of  an 

asceiidiiiL'-  \oltaie  current,  the  ed'eet  iiia.x  lie  to  stimulate  e\piratioi; 
rather  than  inspiration.  Such  results  wdiild  seem  to  indicate  tliat  the 
\a'.ji|s  ciiiMins  hm  l.niils  of  ,ilhr,,il  fih,  ,:•.  hi  II,,  ,■,  si,',i;ili,rii  ,,iil,i\  oiic 
kind  stiiiinlatiiii;  i.ispirat  ion,  the  other,  stimiilatiii;:  expiiation. 

Siipposiiii:   that    siii-h    tillers   exist,    the    next    iplesliou    is,   liow    do   tlie.\ 

'' 'Ill''  stimulate(|    ;it    their   teriiiiiial  ioiis   in   the   liiiiLrs.'     The   most    nal 

iiial   as-.iimption    is   that    llu'    iiieidianical   distention    and   collapse   of  the 
aUeoli  Hhieh  occurs  with  each   respiratory  act,  serves  as  the  stimidus 
an  hypothesis  to  which  sn|ipuit   is  otVered  liy  the  oliservation  that,  wiien 
air  is  lihiwii   intii  the   Iuhlis  so  as  to  distend   the  alvi'oli,  the  animal   iiii 
iiiediatelv    iinikes  a    iniced   expiratory   movement,   whereas   when   the  aii 
IS  siii'ked  out,  the  thorax   assumes  the   iiispirator.v    p<isitioii, 

"I'  'he  iiianv  methods  lliat  iiave  liei'li  employed  to  produce  (listen 
tion  of  the  alveoli,  the  hest  is  uiidoiilitcdly  that  reeeiitl.v  employed  liv 
lialdane  and  Moothliv.'  'I'lie  peistm  or  animal  is  iiiaile  to  respire  Ihroiiu'li 
"  'll''<'  in  whiidi  is  iiiserled  a  three-wa.v  stopeo(d<.  which  communicate^ 
li'her   wiih   III,,  (luiside  air  m-  with   a   side-tuhe   leadiii"-  to  a   s))irometei 

'"■  ''''I'-"  itaininu  air  under  sli'iht    pri'ssiire,  s..  that    when  the  stopeoci 

is  turiieil  hreathiiiL;  lakes  plai'c  auaiiisl  a  (h'tinite  positive  ))ressiire 
!^ii''h  a  milhod  is  uliv  i.iusly  much  more  physioloiiii  than  one  in  wliieli 
the  aii-tulii'  is  suddenly  clamped  at  Mie  end  of  inspiration  and  the  liiiiirs 
left   in  a  distended  condition. 

The  teiiii   iiscil   III  dcsiijnate  ihe  cessation  nf  hreat  hinL'  is  called  iiiiitm 
11 xteiit   to  which   it   oeciiis  Varies  very   eousiderahl.v    in  ilitVereiit   an 


'I'll!    <  (isiKiii.  nr    rni    i;i:>i'ii;  \riii\ 


, mills  iiilil     ill    till'   .-ii^r   III'   liiiili.   ill   (jirri'i-ciil    iiiili\  idllMls.      'I'liiis.    when    :\ 

mall   is  iiiinli'  sudilciily  tn  liii'iitlii'  iiili ni|ii'('sso<l  iiir,  tlic  apiicfi   (iftcii 

hisfs  I'll!-  ;it(cpiit  liiilt'  a  iiiiiiiiti',  llic  |)aHsi'  liciii<r  tlicii  lirnkfii  liy  n  dvv\)  cx- 
piraticpii  I'lilliiwi'il  liy  a  t'lutliiT  paiisi'.  thru  ayaiii  an  cxpirafioii,  and  so 
.111  «illi  pi-ii";i('ssi\  cly  slim-li'i'  panics.  I  )isi'('<;ari|inLr  fur  tlic  |ii'i'sont 
,iiiy  inlliii'iici's  wliic'li  clianui's  in  tin-  coiiipcpsitiun  dI'  iIm'  aii'  in  tiic  liint'v 
ui-  (pf  till'  yascs  in  till'  liiiKPil  iiiiulit  lia\i'  in  piipilncimr  the  aiinca.  wo  may 
i-iiiisiilcr  the  piissilpility  tliat  it  is  tiic  icsiilt  ipI'  alTncnt  tiluMs  in  tin' 
\auiis.  Tliis  is  an  nld  view,  lint  tin'  ninst  recent  experimental  evidence 
lines  lupt  lend  suppciit  tip  it.  It  Was  show  n  hy  Montlilpy  and  lien'y."  I'oi' 
I  \ample,  tiiat  a  similar  apnea,  tlioilirli  indeed  n)'  slmfter  duiati<pn,  could 
iie  pfddlleed  in  duiis  in  wliieli  tlie  pMlmipnai>  Inaiielies  nt'  liotli  vafjus 
!iel\es  liad  lieeii  severed  t>Mp  mniitlis  |)reviiiu>ly.  Tile  apin'a  is.  there- 
fiii-e.  not  a  retle\  of  tlie  vaulis.  am!  must  lie  interpreted  as  due  to  iierv- 
iiils  imi)iilses  i)assini;  to  tlie  respiratory  I'enter  from  sipme  ottier  jtait  ol' 
Mil-  nervous  system,  jierln.ps  from  centers  liiulii'r  np,  or  to  stimuli  trans- 
mitted to  the  respiratory  center  possibly  tliroiiuli  alVereiit  liliers  in  the 
! "spiiatory  muscles. 

The  fornierl\  xcry  pipimlar  llicorv  that  respiration  is  controlled  au- 
;iiiiiaticall.\-  li>'  altcinate  distention  and  collapse  of  the  ahcoli,  actinu: 
'lirouirh  the  atl'i'reiit  fillers  of  the  vauMis  nerve  on  the  respiratory  center 
111  such  a  way  as  to  lirinir  the  o](posite  act  with  each  expiration  and 
inspiration,  must,  theret'ore,  lie  alpaiidoiied.  Hut  \\  i'  can  not  deny  that 
'he  vajrus  ])lays  a  most  important  role  in  the  control  ipf  the  function  of 
'he  respiratory  center.  I'or  apart    from   the  elTect    v.hich   vvc  have  seen  to 

iillipw-  the  severence  of  cipiitiniiity  of  the  nerve,  there  is  the  important 
■  liservatiou   of  Ah-ock  and  others'     that    when    iionpol.-iri/alile  electrodes 

111'  jilaced   oil   the   vayiis   nerve  and   eoiinected    with   a    iralvanoincter,   a 

I'lrrcnt   of  action    nrs   toward    the   end    of  eai-h    iiis])iialioii    in   (|uie1 

iiieatl  ,    i.' :    and    when    the    respirations    are    foii'cd.    a    current    of   action 

i|ipeai's  (iuriiifi:  liotli  inspiration  aiid  expiration.  .\notlier  reason  for 
ill  lievinj;  that  the  vajji  li.-ive  some  important  fiiiiciinu  to  pert'orm  in  con- 
'M'.'tiipu  with  the  control  of  r.'s|iiT'atioii  iN  the  fact,  oliscrved  hy  !•'.  II.  Sciptt," 
■'lilt  in  an  intact  aiiimiil.  vvln'ii  iitmos|iliei'e>  coiilainiii'.;'  incri'asinii'  pi'rccnt- 

iL;es   of   carlioii    dioxide    are    respired,    the    respil'ill  ions    liecome    liotll    deepel' 

iiid  ipiicker,  wliereiis  in  one  whose  v;iui  liiive  lieeii  cut   the  carlion  diox- 

■le  causes  oul.v   a    deepeninu:  of  the    respirat  ions.      I'mm    this     result     it 

ould  a))pi'ar  that  tlii'  Vin;i  exert  jin  iiitliieiice  on  the  rate  of  the  respiiii- 

'ioiis  hut   not   on   their  del)tli.   this  efl'i'ct,  iis  we  shall   see   Ijitcr.  Iieinji'  de- 

"iident  primaril.v  on  chanues  in  the  composition  of  the  liloipil  sui>])l.vina 
■hi-  !-,-s|>i,-ator,v  eentei-.  |l  is  jonhalile  fli.it  l.otli  conli ulliiiu:  .".Ltrrirics  ;icl 
'■luelher.   the   one  serviiiii    to    iiiiiintain    the  eeuter   in    ii    pioper   state   of 


ill 


ii/  :i 


1.  ■•'  : 


': 


\l 


■.\:\\ 


Till    i(i>ni(\Tiii\ 


(■\citlil)ilil\  ,  Jiiid  liciim  ;icli\<'  Id  ;i  u-iiMicr  di'  loss  I'Xtciil  all  liic  lliii' 
while  the  cillitT  iifis  iiliiy  iici';isi(iii;ill>  nil  tllc  •■tuiicd  ill)"  I'L'lltcr.  Tin  i  ■ 
is,  (if  I'oiiisc,  no  (loiilit  tliiit  it  is  lliidiiuli  the  nerves  that  tlie  (ieeasii>ii,i 
aiteraticins  of  iesj)irati(in  (leeiir.  They  a|i|iear  alsn  to  lia\e  a  ecitai 
iiilliieiiee  on  tiic  rliytliiii,  lor  Stewart,  I'ike  aii<l  ( iiithiie''  ohseiveil  tlia' 
al'ter  rcsuseitalion  I'roiii  aeiite  brain  anemia,  tlie  res|iiiatioiis  ulieii  the 
returned  were  of  the  same  rliythiii  as  that  of  the  aililieial  respiiatioi  . 
emi>lo,\ed  duiiiiir  the  resiiseitation. 

Tile   usually   aeeei)le([    jiyimthesis   as    lo   ijie    lueehanisin    jiy    \\  hieli   lli 
iier\f    iiiii)iilses   liasten    the    ies|>iiatoiy    iiin\ cnieiils   is    that   an    allVreii 
iui))iilse  is  transmitted  to  the  lespiraloiy  eeiiter  tnwards  the  end  of  eael 
ins|iiiatioii,   whieli   lias  the  effect   of  inhihilim;  the   inspiiatniy   disehari; 
from  the  eelitei-  and  thus  eiittinu'  slmit  the  act   of  iiis|>ii-a1inn  so  that  (a 
l>ira1inn    aiitimiatieally   supeix  eiies.      'I'liis   explanation    is    in    ajiieeuieii' 
with  tlie   faet  that  (piiet    inspiration   iinoUcs  aetivit.\    mi  the  l>art  of  tie 
respiratory    mnseles,   whereas  e\pirali(.n    is   usually  almost    entirely   |>a^ 
sivc    heinir  due  to  tlie  return  to  a   resliiii;  position  of  the  stretched  :iimI 
o,   ,ilaced   structures.     ( »n   the  other  hand,   in    forced  resiiiratioii    and    ii 
ecita'ii  animals  under  normal  conditions,  expiration  hecoiues  aeti\e,  ii: 
which  exi'iit  a  eiirrent  of  action  hecoiiies  e\ideiii   in  the  \auus  n  mm'  il;ii 
iiiir  the  ex))irator.v  plinse. 

The  superior  larynjre;il  liraiieh  of  the  xaijiis  should  really  be  elassilicil 
as  one  of  tliose  nerves  that  ha\e  an  occasional  inlliieiico  on  the  respiratoi  \ 
center,  its  pnrtienlar  function  lieiim-  in  connection  with  the  act  of  eoiisjh 

ill!.'.    When  a  foreiyn  Iiody  irritates  tlio  im ns  memhrane  of  the  larynx 

the  nerv(>  fihers  traii--mit  impulsos  to  tlie  lespiratory  center  wliicli  ex 
<'ite  a  violent  expiration  and  at  the  same  timi>  cause  the  vlotlis  to  elosiv 
The  closure  of  the  jilottis  lasts,  however,  only  durinir  the  first  part  of 
the  expiration:  it  then  opens,  with  the  result  that  the  sudden  release  oi 
intrapiiliiionic  pressure  causes  the  expulsion  of  the  t'oreiyn  sulistanc 
in  the  air  pnssajios. 


ClIAI'TKi;  XXX\  III 


TIIK  ((iNTIJOI.  (»I'  HKsi'lli.VTKtN   .Cmt'd) 
THE  HORMONE  CONTROL  OF  THE  RESPIRATORY  CENTER 


^ 


•Just  as  tlic  ili\lliiiiifal  ;uti\ity  of  tlie  lioart  is  readily  iullueiicctl  1)\- 
■  liaiisrcs  ill  the  ciiiiiiiiisiiion  of  the  IiIimmI  siipplyiiii;  it,  so  also  is  tliat  of 
ihc  rcspiratoiy  I'cuiir.  In  tlii'  case  ol'  tlic  iicait  it  is  IIk'  cations  -cal- 
■iiiin,   potassium    and   soiliiini   -tliat    liavo    tin;    most    )ii(mouiiccd    cfTi'ct. 


>li( 


icreas  iii  tin'  rase  ot  tlic  rcspna 


ftli 


i-y  center  it  i>  lai'iri'ly  tiie  relative  con- 
ion    concentiation 


entration    of    Iiydrniicn    ami    li.\«lro\yl    ions     the    II 

<'m^   of  the  lilood.     Tliis  inllucMi'c  can  lie  shown   in  a  ironoral  ■w  a.\    In 

iMJecfinft  acid  of  alkaline  snlntiiu's  intu  the  i)eiii>heial  end  of  the  carotid 


lei-v  of  an   anes 


thet 


i/el   anin;a 


1,  or  hetter  still  of  one  that  lias  li 


I'ceiTliratcd.  Acid  in.jectinns  stimnlate  the  resi>ii-ator>-  activity,  alka- 
nc  injections  tend  to  depress  it.  When  the  acid  ur  alkaline  solutinns 
re  injected  intia\  enonsly  in  other  parts  of  the  hody.  so  that  tlu'v  lie- 
iine  thoroiiirhly  mixed 
'ached,  the  effects  are 


ith  the  1.1 1  liefore  tl 


not  nearly  so  jn-nn 


le  respiratory  center  is 
onnced.  because  the  hiiffer  in- 


■''lence  of  the  lihHid  has  tiiiie  tn  develop   i  see  p;|t;c  '.'M) . 

From  the  results  ul'  such  injection  experiments,  liowever,  one  could 
'tilt  draw  tlie  concliisicin  that  under  iu)nii'ii  conditions  the  activity  of 
'he  resi)irator\'  center  is  alTecled  liy  iiieasurahle  chanires  in  (',,  of  tlie 
Mood,  for,  as  we  have  seen,  constancy  of  (",,  is  one  ot'  tlie  most  rcniark- 
thle  properties  of  the  animal  tliiids.  To  justify  the  conchision  that  the 
^ 'spiratory  center  is  alTectciI  hy  clianires  in  (',,,  it  is  neeosar.v  to  oliserve 
*'ie  behavior  of  sunie  easily  iiieasni'able  iicid  nv  alkaline  constituent  of 
'he  blood  that  nnderj;oes  chauires  in  amount  that  are  i)ro])cirtional  to  an 


UiTation    in    (' 


In 


der    t 


o    niHlci'stand    what    this    acid    or    basic 


ibstanee  may  he.  it  will  be  adxisable  to  recapitulate  the  main  factors 
iiiiirned  iti  inaintainin<r  <'n  at  a  constant  level.  This  value  is  obviously 
'  I'enib'nt  ui)on  the  Jialanee  between  basic  nml  jicid  substances,  so  tbat 
M\-   variations  which   it  underjioes  must   b(»  caused  bv  chanures  in  the 


ativc  amount    of  one   of  tliese.     Chanires   in   linsr   mav   oc 


cur.   exofre- 


■nslv.  bv  alterintr  t 


le  alKali  eon 


tent  of  the  food. 


or.  emloirenouslv.  in 


oils  ways  but  ])ar1i<'ularl>-  liy  variations  in  the  amount  of  animonia 


iliiced  (luriii}!;  tin 


if  liieiabc>ii->i 


11  (if  lU'otein. 


Th 


us.  wlieu  su( 


demands  are  made  b\-  the  oriraiiism  for  an  increased  amount  of  base 


'    J' 


1  ' 


.?.x- 


1 1  p 


:i:i(i 


■nil     l!l  >l'll!\lln\ 


1 1 
1  f 


M. 


r      i 


W 


i:    ; , 

I-  i^ 


l|.:: 


ii-! 


the  :iiiiiii(i  uiiiii|is     -,|pllt    iiiT   iidiii    till'  iiniiihi   ImkIIi's     I nii^   (•iiii\('iti''l 

into  .'iiMiiioniii  iiislciiil  111'  into  ilic  ni'iiliiil  siilisliiiici'.  iifcn.  Miit  tlic  cliiri 
\.iii;ilii)hs  si'ciii  t.i  i-iiiicirii  iiiiih  r-Mtlicr  tluiii  llii'  liii>ii'  >iilistjiii<M's.  Tlics'' 
:iciiK  iii;i.\  Im'  (||\ii|('i|  iiilii  tliicc  ui-uiips:  /('./((/  innriiiiinr  'iriih.  I'fprcsciifcil 
liv  iiliiis|)lMiiic;  fiiiil  iiri/iniic  ni  ills,  rcpri'scnlcd  liy  luetic;  iiMd  riilnlih 
mills,  i-cpn'sriit.'.!  I.\  .•MliiHii  (Iiii\i(lc.  of  tlirsi'  tlllec  un.ups,  till'  (irsi 
nIkiHs  till'  least  teiideiiey  In  eliauiic,  and  tiie  tllild,  tlie  u:i('Mtest.  Cliallfies 
in  tlie  seedinl  i.'i(inp  ili\ed  .ili;aiii<'  aeid'^'  are  ell'ecte.l  jjaitly  liy  excrctinii 
tliKiniili  til.'  mine  ami  partly  liy  Dxidalidii  into  volatile  ai'id.  The  snd 
den  and  rapid  eliaiiL'es  in  tlie  tliird  irrmip  are  liroii;.dit  alioiit  !(>•  tlio  dif 
I'llsioii   ol'  tlie  ('(»,  ol'  tlie  Mood   into  tlie  aheolai'  air.     (iioss  eliantres  in 

tlie  acid   eontent   of  tlie   111 1   are   lliefel'ore   mainly   elVeeted    tlirmiirli   al 

teratioii  ill  the  e\eretion  of  llie  lixeil  aeids.  whereas  siiclden  ehallires  are 
ell'eeted  li\  excretion  of  tile  volatili'  aciil.  It  is  important  to  note  hen 
that  the  li\ed  oi-eaiiic  acids  do  not  |iarticipate  to  an.\-  ureat  extent  in 
the  malvciip  of  the  aciil  content  of  noriiial  Mood:  they  appear  only  niidei 
niiiisiial  conditions,  ms  in  dyspnea.  The  variations  in  ('„  tliiil  ordinarily 
alVect  the  actixity  of  the  respiratory  center  are  therefore  de]>on(leni 
ii|ion  ehaii^zes  in  the  volatih'  acid,  a  dir.'ct  measure  of  wliicdi  is  fonml 
in  the  ti-nsion  of  (  (  i  in  the  liloo.l.  The  corivlation  hetwecii  ('„  of  the 
liloiid  and  respiratory  acli\ity  must  hi'  a  \ery  close  one  if  ('n  is  to  he 
iiiaintaincd. 

The  Laws  of  Gases.  In  order  to  understand  he  inineiples  ni>oii  which 
alterations  in  < '< »  tension  are  di'pciident,  it  will  i-e  necessary  for  ns  t" 
review  hrielly  some  of  //,,  i/ns  Imi's.  Amoliy'  these  li-ws  llie  first  in  iiii 
portaiiee  ']<  the  /(/((■  (//'  /in  ssiin .  which  states  tha*.  oiiiei'  tliin<rs  lieiicj 
eipial.  the  pressure  of  a  uas  is  iinersely  ])roport ional  to  its  \dliiino:  i: 
a  yas  occiipyinu-  a  certain  \oliiiiii'  is  compressed  liy  a  piiniji  so  that  it  oc 
eiipies  one  half  of  its  pre\  ions  volume,  its  ))ressure  will  liecome  doiiMed 
The  second  is  the  Imr  of  iiiirlliil  i,rissi(ri,  \\\\u-h  stales  that  the  partial 
pressure  of  a  '.ias  in  a  mixture  of  trases,  Inivinu'-  no  ai'tioii  on  one  another 
is  eipial  to  that  which  ilds  particular  yas  w.udd  exert  did  it  .iloiio  oc 
ciipy  the  space  occupied  liy  the  mixture.  Thus,  almos|)herie  air  consist- 
rouuhly  of  7!'  \oluiiies  jier  I'l'iit  of  nitroiieii  and  -]    of  oxyircn  :  the  jiar 

■'1 

tial    liiessiire    of    the    oxvueli    is    therel'ore    eiiiial    to   ~       '    "''"  linn.       1 1 1^ 

'  Kill 

lliis   last    Ijeiiie  heine    the  liar trie   pressure  of  air  al    sea    level.      Tli' 

third  is  the  law  of  si, hit  inn  nf  i/ii.si  s,  which   is  to  the  elVcct   that  the  amoiili' 

of  u-as  which  eiies  into  solution  in  a  liipiid  lia\iiiir  no  chemical  attractioi 

for  the  iras,  is  proportional   to  the  jiartial   inessiire  of  sras.      If  water  i- 

e\piised  to  ail',  the  amount   of  o\yui'n   which  it  dissol\cs  will  he  l!ie  saiii- 

as  if  the  water  had  hcen  exposed  to  ox\  L'eii  at   a   pressure  eipial  to  tlnr 


t;ii;  nivi'ifni.  (IK   iin:  ursi'invrinv 


fia? 


ii    tlic   piiiliiil    iiicsNiiii'   wliicli    il    |ii(m1ui'<'s   ill   iiir.      Tlic   siiiiii'   will   !»' 

Ill'  ciisc  willi  (lie  iiilriii.'rii  ul'  llic  iiir.     Tlic  iii-tiial  iiiiinlliit  nl'  Kiis  wliicli 

i'li-oiiics  (lissnlvril  ill  till'  lliiiil,  jircssiirt'  iiiul  li'iiipci-iitiiic  liciiiy:  fimstniit, 

li'ix'iiils  jiiirllx  iiii  tlic  iiiitiiic  III'  llic  ifiis  iinil  ))iirtly  mi  tlic  luitiirc  <if  tlic 

iliijil.     Fur  cxiiiii|ilc,  tlic  sdliiliility  oi"  (ixy<„'cii   in   wfifci-  is  cniisiilcriiMv 

lill'ci-ciit  fniiii  tli;it  in  ii  iiciitnil  nil:  nr,  t,ikin<.'  tlic  siinio  solvent,  iiitro- 

■rii  iiiul  CO.,  (Ill  iiiit  ijissdlvc  to  llic  siiiiic  extent   in  wiitcr.     It  liccnnics 

iii'cssiiry.  tlicr<'f'oic,   in   e;i!eul!itiiii,'  wliiit    jiiiioniit    of  n   ]iiirticuljir   <riis 

\\ill  (lissiilvc  ill  il   piirticiitiir  tliiiil  to  use  ii   tijriire  kiinwii  as  tlio  rm  iliriciil 

nf  sohihllihi  of  tlie  '4:fis     tliiit  is,  tlic  iiiiioiiiit   nt'  "iiis  tiikcii  up  liy  a  unit 

Illume  III'  lliiiil  at  staiiilaiil  teiiipcratui'c  ami  )ircssure:  I'm'  cxamiile.  to 

,i\    that   tlic  eoenicient  ot'  alismptiiiii   of  nitinu'cii   in   water  at  0"   <'.  is 

iiirj:!!*  iiieaiis  tliat.  at   tills  teiiipcialuie  and   at    normal   liaroiiictric   ]ircs- 

>iiic.  1  c.c.  lit'  water  will  ilissulve  (l.()"J:'.!t  e.e.  of  nit niircii  wlicn  exi)osp(l  fn 

pure  atiiios|)liere  of  tliis  iras.     (»li\iously,  tlien,  if  water  w  cic  exixiseil 

I  7!t  per  cent  of  an  atmosplicre  nf  iiitro^'cn   i  as  in  ain  tlie  aiiioiint  wliieli 

70 


Iiceome  ilissolvcil  in  caeli  e.e.   wonlil 


11)11 


0.02:W  ^-  0.0180  CO. 


In  solutions  eontainiiiir  no  elicmieal  sulistanees  witli  wliieli  tlio  iras  can 
liter  into  (•oiiiliinatioii.   it   is  cviileiit   that   tlic  teiisinii   of  tlie  ;ras  will   be 

■  iMportional  to  the  aiiiount   of  «ras  that   can   he  displaced  or  piiiiiped  out 
•iiii  the  Hliid.     On  the  other  hand,  when  a  eheinical  coiii]i()uiid  is  forniod. 

•lie  eoiiiliiiicd  fras  will  exercise  no  direct  intluciice  on  the  tciisinn.  so  that 
•liis  will  he  indciicndcnt  of  the  amount :  in  niirli  rnxra  .iipnrafi  uirthixh 
rill  Uiiv(  to  hi  iisiil  fur  ilii  ill  II  niniititiiiii  nf  iimiiKiit  niiil  Ini.siuii.  T<ot 
,>  lake  the  i-asc  of  |)iire  water  cxiioscil  to  an  atniosphere  of  CO^:  the 
iiiiiinit  of  CO,  which  jiocs  into  solution  will  deiicr.d  entirely  on  tlio 
■M'ssure.  Tf  a  trace  of  alkali  is  dissolved  in  the  water,  however,  some 
■<'  the  CO.  will  liecoiiic  comhined  to  t'oini  earhnliate,  so  that  a  much 
iiiu'cr  i|iian1i1y  of  CO,  will  lie  disjilaccalilc  from  the  solution  (as  liy 
'liliiuj:  a  mineral  acid  to  it  i  than  corresiionds  to  the  tension  of  CO_.  in 
'ir  atmosphere  suironndiinr  it.  Since  lilood  contains  alkali  the  coiid' 
^'iiis  are  analoirous  with  those  of  a  weak  alkaline  solution. 

The  Tension  of  CO,  and  0,  in  the  Arterial  Blood.— If  we  were  to 
iss  hhiod  at  liody  tcmi)crature  in  a  very  thin  film  (i\cr  the  walls  of  a 
■  nfined  sjiace  coiitainiiifi  a  mixture  of  irascs  one  of  which  was  CO.,  it 

■  i\  idcnt  that  the  pcreentajic  of  CO,  in  the  atiiiosplipre  eontainnl  in 
'  is  space  Would  rciiiain  iinchaiiircd  only  when  the  tension  of  this  iras  in 

•  r  lilood  was  the  same  as  that  in  the  contincd  atiiios])hero.  If.  on  the 
ilierliand,  the  tension  of  CO,  in  the  lilood  slnnild  c(irrcs|)on(l  to  a  per- 
■iitajre  that  is  hiirlici'  than  that  in  the  atniosphere,  then  CO^  would  ilif- 
i -e  from  the  lilood.  and  at  the  end  of  the  experiment  an  analysis  of  the 


i-i 


ii  iii 


;i"]f< 


Tin:  Ki>i'iR.\Tiii\ 


i    i  '< 

1 

!' 

i 

'f 


JllliK.splific  ill  till'  Npn.-r  w.iiiM  slii.u  Hull  llic  CO.  pcivi'iitiiHf  liad  Immi. 
liiis.'.l.  If  the  Mood  coiiliiin,.,!  a  lower  tension  tliaii  that  eorrespondinii 
to  the  per.-elitasre  of  ('(»,  in  the  sp.iee.  some  of  tht"  CO,  wonld  difTlis. 
into  the  Idood.  and  its  p.iveiita«e  in  tlie  alinosph.'ie  would  he  lowered 
I5v  sueeessively  eN|M.siiii;  Mood  |,,  ujis  mixtures  that  eontain  sli-rlitl.v 
ditVerent  pereeiita«es  of  CO.,  «e  shoiiM  ultimately  lind  one  with  which 
the  free  CO,  in  the  lilood  wns  in  p.-rfeet  e.|nililniiini.  and  wc  should  hr 
aide  to  state  that  the  tension  of  this  u'as  in  the  hiood  was  ef|iial  to  a 
eertain  lu-rei'iitau.'  in  the  atmospluMv  surroiindinj;  the  hlood  (see  I-'i--' 
121  1. 

AFaiiy  forms  of  apparatus  liased  on  the  ahove  priiieiple  have  been  in 

^ '■"*''''    •'"'■   'I \aiiiinatioii   of  the   tension   of   the   leases   in   tlii>   1iIo<hI 

The  most   aeeiiralr   is  that   devised   l,y    Krojih."*  the   luineiple   of  wliii-h 


CO      **•»•<"» 


■'--^-"' 


■^^'^'I'^i- 


!} 


'-^''■^  ■^^-^/'■'/^Bggy^.^^^^^^-^ri^J^^.^^.^v'. 


•■*     t  i 


CO,     i  n  »'  »wt 


11 


vv     |,nti,„.I,-     f.. ,„,,:,., r,.-:.nl    „t     tin-     t.  „-,..„     ,,1     eO..     i„     ].],.„].       '1  1.. 
"_    M.-i.,n    ,.f    l,|u...|    1^    M,].|.,,«..|    tu    li>-    5.75. 


<litrers  sliirhtly  from  that  just  deseril.ed  in  that  a  l.iiM.lo  of  air  is 
exposed  to  a  relatively  lar-e  .|uantity  of  l.lood,  so  that  after  a  time 
aetual  e.|uili:.rium  of  -;,s  tension  heeomes  estal.lished  l.etween  the  huli 
Me  and  the  -ases  of  the  Mood.  This  apparatus  is  shown  in  Fifis.  lL>-.> 
and  ]•_>:!.  It  eonsists  of  a  graduated  tuhe  of  narrow  Ix.re  sur- 
rounded l,y  a  water  jaeket.  To  the  upi"'i'  ''nd  of  the  graduated  tulie 
a  small  syrin-e  is  attached.  The  lowr  end  of  the  Graduated  tul.o  ex 
pands  int..  a  thistle-shaped  hull.,  closed  l.elow  I.y  a  eork.  thn.u-rh  which 
IS  inserted  a  tul.e  (inflow  tul.ei  endintr  near  the  to).  ..f  the  bull,  in  a 
fine  openiiifT  and  eoiinecte.l  outside  with  an  artery.  An  outllow  tul.e  is 
also  eoiinecte.l  with  the  tliistle-shaped  Lull.. 

At  the  l.efriunin-  of  the  experiment  the  thistle-shaped  hull,  and  the 
irraduated  tul.e  are  filled  with  physiologic  saline.  ]5v  means  of  the 
syrinpre  a  small  l.ul.l.le  of  air  is  then   intr..dueed,  so  that  it  lies  at  tlic 


Tin:    ciiNTIinl.   Ill'   •JMi:    lil'.sl'llc  M'|ii\ 


;j;{!« 


.11 


llirtinn  III'  till'  I  lli^l  li'  sliiiiMil   liillli  Illnl  llii'  <j;l'ili|llilli'i|  tllln'.      As  till'  lilmiil 


i>  iiliiiwt'il  til  ciitcj-  tliriiii^li  till'  iiiliiiw  tiilii'.  It  IS  e.jci-tfil  in  ii  lini*  stiriiiii 
nroiiml  tlio  liiililili"  III'  nil',  V  liicli  iimvi's  iilimit  in  tlic  sti'i'iiiii.  Tlit-  liloiiil 
■  lispliirrs  tlic  saliiii'  uiit  iil'  tin-  liiilli  iiilii  tlii'  siiir  tiilio.  AI'tiT  the  liilb- 
Mc  liiis  lii'i'ii  siili.ji'i'tiij  til  till'  inlliii'iii'c  III'  till'  liliinil  I'lir  siiiiii-  iiilmiti'*. 
;lir  U'iisi's  in  it  I'lnnr  ilitn  I'l  ij'crt  ri|llilllil'illin  witli  tlinsc  in  tlic  Illiiml. 
Till'  |ui('i'iil;iLri'  111  (».  iiiiil  III,  ill  till'  liiililili'  will  tliiMi'tnro  ('ori'csp'Hiii 
til  till'  Iciisiiiii  111'  tlirsc  triisi's  in  llii'  liliMiil.  Till'  aniilysis  is  crtVctt'd  l»,\' 
'li.iv^iiiLr  till'  linlililr  intii  llir  urjiduiili'il  Inlii'  liy   iiiriins  nt'  the  syringe. 


I'l;    I ' 


i-'ii.-,  1' 


li^r.    I_'..'       Tin     Kit-    .lii;i]>>i'«    |.ipitt.'    (■■]    ttl'     r.ii' i..t,.ii,-,p.  l<  I    -Iim«  n    ni    J-'n^.     \j.i,       l'"..i     ,|,  ,,  ,  i|.t,,,n 
.i.rit<-\t,      (l-'nim    A.    KroKli.l 

l'ii(.    KM       Miiri.t.iii"i1uii  r.    I..   In     in-.n..l    :t:l..   :i    M 1    \.--.I       Tli.     -n   ,11    cinl,-    r<|.ri~iiu-    lli. 

'■"!■    "t   air.      l'"'.i-    Iiirth.T   il.-,i  i|.li.  !i    ^(  «■   (.'i.ttM.      I  I'r  'iii    .\.    Kt..i;li.i 

Mitasiiiini:  its  capacity,  ti'aiisl'cirinir  it  into  a  luill)  cuiitaininsi:  KOlI, 
.  hii'li  alisiiili.s  tlic  CO..  tlicii  takiiiir  it  hacU  iiitu  tlic  capillary  tulip  anil 
.iirain  measuring:.  The  sliiinkairc  uiivionsly  cnircsiiniKis  to  tlic  amount 
"!'  CO^.  The  Inilihic  is  then  traiisreiieil  into  iintassiniii  pyroijallate  solu- 
'ioii.  where  the  O^  is  alisoilinl.* 

The  Tension  of  CO..  and  0,  in  Alveolar  Air.     Havinu:  seen  liow  we 
may  (Icterniinc  the  lension  ol'  the  trascs  in  Mood,  \\c  must  now  coiisiilcf 

'SitK-c  iha  ;i1m.v.-  w.i~  \m.ii.ii.  a  iu..r.'   .IVniMU    1..11..1111  l,  i    .!.m.,,I    1,v    iIi.    I.iI.    '1  .    C,     II1...I11    lia- 
■11  •l.s.ril,,,!  I,y    (rSulliv.iil    (  ^lll    .L.iir     l'by.i..l.   S.l  t  .    I'Hsi. 


H 


!  '  I 


' .  I  *;.l 


'  I 


I  ii 


|i 


1 


f: 


i  il  :i ' 


1 1  ^  IP 


I  if 


it  ■ 


■I 


I  ,     I 


:M() 


Tin     UrM'IH\TUi\ 


til.'  iiK'tliixl  l.v  wlii.h  III,.  iniNinns  i.f  ilicsf  miscis  ill  iilvci.lar  nir  i-im  \w 
(Iclniiiiiic.l      Tlic  simplest   iiiii!   until   recently  the  iiinsf  acciliiife  iiietliixi 

is  flint  of  llaMaiie  ami  l'iiestle\ ,'  ■    This sisis  in  liax  Ink  nil  individual. 

"itii  liis  nostrils  elaiiiped,  l.reatlie  .|uiell.v  tliiouifli  a  |>i f  hose  pipe 

■■''"""  '■'  I''!'  '"'Il-'-  "lii'li  liiis  at  the  iiiiMith  end  a  short  side-tulie  lead 

iMy  to  an   evaeiiated   !.'as-sainplinjf  liulli  of  aliout    .Ml  e.e.   eapaeily.*      After 

the  sulijeet    has  I me  aeeustomed   to   Kreathiiij;   thloii-ill    the   tube,   h<' 

IS  asked  to  make  a  joreed  expiration  and  at   the  end  of  it  to  close  the 

moiithpieee   with    his   ton-rue.      At   this   meiil    the  oporator  opens   the 

lap   of  the   sampliuir   tui.e,   allow  iuLT   the   air    from   the    tiihiiiK   throud, 
"hieh  the  iiKlividual  has  made  the  loreed  ..xpiratioii  to  rush  in  and  fill 
It.     This  sample  represents  the  air  from  the  alveoli    i  see  pajri"  .■1(12).  and 
is  analyzed  for  p.Teentanes  of  ('(».  and  O,,.      Since  each  iioniial  inspira 
tion  dilutes  the  alveolar  air  somi'«liat,  it   is  iieeessaiy,   for  eonstaiit  le 


l-iK.     l.M.     .\,.,..„a.u.    f.„     ,„ll,...,i„„    ,„•    a    -a.Mi.l.-    ..I    ;,l>,-„!,,r    ai,     l,v     ll;,l,l,„„.-,    „„H,..,|        1,    ,. 
I.illir    Im    n>i'    a    miMllll|.ini-    lliin    a    lii.isk. 

suits,  to  make  two  analyses  of  alveolar  air  from  each  sul.Jeef.  one  taken 
at  the  end  of  a  normal  insi)iiiition  ami  the  other  at  the  end  of  normal 
cvpiration.  The  averaire  of  the  two  results  is  taken  as  the  composition 
of  the  alveolar  air. 

""  " "'"  "'■  "'«■  'lirticiilty   in  seeuriny;  intclli-reni   cooperation   in  the 

apiilicatioM  of  this  in.  ihod,  particularly  with  children,  others  have  heen 
devised.  One  of  the  simplest  is  that  of  Kride;  icia.  which  is  a  modifica 
tiim  of  the  llaldanc-l'ricstley  nictho.l,  the  apparatus  for  whidi  is  shown 
in  the  fiijurc  (Fin.  I2r.i.  and  th..  manipulati,ui  of  which  is  outlined  in 
the  IciTcnd.  Another  is  to  fake  a  mixed  sample  of  the  very  last  portion 
of  several  normal  expirations.  On  account  of  the  cMended  use  which  is 
'"''">''  """'*'  "'■  iii''iisurcments  of  alveolar  air  composifh.n.   lioth    in   lah- 

.1-   lh',."„',''"'rr,',"    ''"    ''■;-  -■■""l'""«  """■  "    '-  "'".1.  ".'...•  ....,v,n„-„    ail.!  ,,|n.,IK    ;,>-,,n,il.-  ,..  ,„ v   ,„„ 


Tin;  itiMKoi,  Df  Tin;  hisiikatihn 


:U] 


I-Mil.l,    tin. 

I   th.-    .iir 

-"hilioii  ,,( 
I  l>ul  »illi 
ill.-  fvtcv* 
111.  II    Mll.inc-i 

'III      I.V.I      ,1 


l'riil.int.i\  .ipi-ai.tlii-.  tur  inia^ininj;  ilit  <'n,  iti  iilv(.>lar  .m.  Tlu-  i-.r^.-n  .  sj.i' 
mil!.  III.  Inl.i-  Ailli  Ihi  ,l..|.c,„  k«  il»  iii  I  A  i-  .:.-..!  .iii.l  llu-  ml..-  |.|.....l  III  Miil.i 
atlci-  whi.li  /*  i-  lii'l.'.l  a-,  iti  II.  Tuc  . 1111,1). |»M  ...I. mm  ..f  a.r  .■'luaN  IMO  »•  .•• 
a»l^llt•  .i!kati  !-  iHiw  >iu  Ivf.l  11. 1. 1  (■  with  st..i>f...  k^  ..»  in  11.  /*  1*  tlicn  tiiriif.l  a> 
I  -nil  .li.M.I.  an. I  th.  alkali  ...lull. .11  alL.iu.l  I.,  .iilfi  /■.  ..ft.  1  wliuh  II  n  Inriif.l  . 
.f  alk.ill  •■..lull. Ill  III  I  all. .«.■.!  Ii.  mil  ..ul  .unl  th.  l.;i..ll.  -Iial..  .1.  Th.-  hun-lt.' 
-.'I  111'  !■•  II  III  .  .sln.l.r  iif  «at.i.  «ith  H  a^  in  III  Aft.r  all..«iiii!  I.t  .....|ii 
uhi.  h    tin-    V,  lUr    -t  in.l-    ciM  .    tin     n.  r    i.  ill    ..t    l'<  ' 


tkj 

tea) 

i<ty        %  % 

— ! 

V 

;        i/.  llLt^urfJ 

la 

/ 

-. 

17 

) 

y 

16 
IS 

■ 

1 

.  \ ' 

.1 

~1  ' 

rt 

■ 

\ 

13 

■ 

\ 

m 

■ 

V 

II 

1 

10 

(W 

0, 

i 

1 

♦ 

.  0— 

COj 

a 

.      ■     ■      - 

^~^^^-:-^ 

•1 

/ 

30^  to' 

.iO        ^          t9         XO 

.10      «/      J»      J       w 

I'lK.  lJf>.— Curves  tn  >hi.\v  the  rt  latini;s,hij,  hiiwicn  tlic  *>^  an.i  Ct  ^.  itiisi-.iis  in  alveolar  air 
'!"it(M|  Itiif.)  and  arti-rial  M.toil  (lontirm.ms  liius).  It  will  l.e  (iltM-rvnl  that  the  tension  nf  TO;. 
>  M'mmI  is  >itKlitIy  alnivr  that  in  .ilv<nl;ir  .irr.  tint  that  the  fversc  rt  l;itinii^hi|i  ohtains  (or  i\-  \\\ 
fie  np[)er  part  of  the  curve  the  O-  tension  ju  the  alveolar  air  wa-  t  vperiinentally  alleieil.  causinn 
<i>rrrspnn.iiii(t  alteration  in  tlu-  <  Jj  ten^inti  uf  the  hloiMJ,  Thi'<  n-'iill  is  i.f  pra.  tiial  slRnituaiiri 
''     tniniectiun     with     U.     all<'r.-ili>>li>     in     uns     nnisiiKiniT      1. II.  11111. iiii.i       .1.'  il''i..nt       \       :ii>.l      \f        k'r..i7li    I 


till        \i\         till.         Illliini.  lII|->        ll'IMI         !■•       Ill         |l|<IVIIl>ll        ^I^IMMIillMl 

pnisui-iny.    piu  iiinoiiia.    it'.'.      (I'lom    A.    and    M.    Ktogh  j 


M  Ii 


$ 


I 

-   '^1 

! 

■     ,-* 

^1 

f 

-f 

I  m 


1 

i   111 


I  >i  t 


•UL' 


rm    Ki^i'iiivTiiiv 


'     H 


r  :1! 


k^H 


i 


hl^ 


hi 

1 

i:     '  i 

I. niton    iiri>l   in   rlinii-iil    \\><\U.  ji   >|in-i,il   .•li.ipl.r    li.is   Ihtm   iliMitcil   tii   lli. 
sul.j.Tt,  jfiviiit:  ill  il.'tiiil  till'  iiii>r<'  i- nt  iiirl  lin.N  ,|,.\  iv-d  l.y  It.  (i.  I'l-iiirr 

l.iiNtly,  it  sli.Hil.l  he  Miiti'il  tliiit  MViijil  oI.mmmts  t)clit'V<'  that  a  hum. 
ii'liiilili'  •■stiiiialc  Ml'  til,,  alv  I'dlar  I.'iivion  i.l'  •'(•  :  aii<l  i>l'  O,  ■  .-an  ]„•  iim<I' 
\'\  aluilv/iiiL'  a  *aiii|il<'  of  cii'diiiai  .\  <  \|iiii'.|  air  ami  filifiilaliin;  tlir  pn 
<Tiitiiu'i'<i  lit'  (■(•  affl  U.  ill  III,.  alM'ulai  aif  l.\  alhn\  iiii.'  a  iMiiistant  .|fai| 
s|iiifO  ('ii|iai'ity  I'l'  no  imv  i|\r(ii»li.  I'tiv'. 

It'  wc  coiiiiiai.'  til.'  CO    li'iisioii   i,r  ait.'iial   M I.  as  iiirasiiiril   l>v   tin' 

l\ro(fli   iii.mIiimI.   \\itli    that   ol'  ah.nhir  air.   « <■  shall   fiii<!    that    th.ic   is  a 
rciiiarUaiilr  niir.'s|M,ii,|.'i ,  iiicliratiinj,  thrii'lorc,  that.  «  hni  the  arterial 


"■    '~"l'l    I'ii^    ill    lln-    i.iv     il;,.    l.ii-k.n   (f    I'd.    i,,    ih,.    ,,U...l;ir    ri     «i 
■  ~r>-"' ■ii'.-.l!y    .,'i.r..|.      iI't.mi,     \     ,i,„l    M,    Kl-.^I,   I 

hliMxl  leaves  the  alv.oli,  its  partial  |iivs>iuv  ,,r  tension  of  ('(»_,  is  e\aetl.\ 
equal  to  that  ill  the  alveolar  air.  This  is  show  i  in  the  aceoiiipaiiviii".; 
eiirvps  of  experinienis  |.ertoiineil  h.v  Kroirh.  The  .lotted  line  in  ihes. 
enrves  represents  the  tension  of  Cd.  ,,r  (t,  in  alvc'olar  air,  and  the  eon- 

tiinions    line,    these    tensions    in    arterial    l.lood.      Clos Tes])ondonee 

uill   he  oliserxed    lietueeii    the  CO     eiirves  even    when   sii.lden   eliailKPS   in 

alveolar  (O     were   iml ,|   l,y   aiiilicial    means.      In   the   i-ase   of  the  O 

tensions,   ho^^e\el■,   that   of  the   Mood   is  always  louei'  than   that   of  tl 
alvpolar  air,  the  dit't'ereiiees  l.eiiiL'  espeeially   marked  when   the  0„  t 
sill!)  in  the  alveoli  is  raised  i  Kitrs.  VJll  and  127  i. 

Tension  of  CO,  in  Venous  Blood.     If  vw  eNamlne  the  CO..  tonsjon  of 
the  vpiions  l.lood  eoiiiiiiju'  to  the  Iniisis,  ue  shall  (ind  that  it  is  distinctlv 


II' 
en 


Tin;    (ONTRilI.   OK    Till     KIM'IHATinV 


3n 


hiu'h'T 


III  tliiil  ill  ilif  iihi'iilar  nil'.     Tlip  rnrlicst  iiicHkmI  fur  iin-iiMiii'intr 


sti'fl  ill  |>ii>siiiu  ;,  linii;  catlicttT  iiit"  tlip  riiflit  lirniichiis  mid  flicii 
Mi\c  tl.'  iipi'ii  cud  of  tlio  riitliffcr  liy  iiifliitiiii;  II 
il'ullii.    Till'  rnicvMil  nl'  nil-  in  tin'  riulit  liiiif;  in  lliorchy 


i|iii'killl{  tlir    |(|ISM1'. 


illiliiT  riilliii'  III'  ^111 


•  I  r\  I'llti 


i! 


saliipli'  III'  t'    ■  stilu'liiilit  iiir  •'( 


II  Iir  riiiiiivfd  iiiid  iiimlw.rd. 


Ill  sui'Ii  a  rasi',  liiiw  ('\  cr,  till-  liliiiid  will  liaM'  riniilatrd  m'M'I'ii!  liiiifs 
miiiid  till"  lindy,  and  with  mily  niic  liiiitr  nprraliiiir  tlii'  iIhU  is  iiiciirrcd 
iliat  iimri'  CO.,  is  ln'iiii.'  diMdiiirifi'd  iiitn  tlif  hlm-kcd  liini;  fliaii  cur- 
I.  >i)iiiiii|s  til  till'  ti'iisiiiii  nt'CIV  iif  M'lHiiis  liliMid  iiiidt'i-  iinriiial  i-oiiditiiiiiH. 
Miifli  iiKiro  i>rai'tii-al  iiit'tlnids  nn'  llmsi'  of  llaldiiiii',  Yiiiidi'li  llrndiM'- 
sMii  and  I{.  (J.  I'cari'c,  wliiidi  firo  iiiin-li  tlii'  satni'  in  |ti-iiiriplc.  In  IViircc's 
imiliiid,  till'  pi'i'soii  lii'st  of  all  inspiri's  from  a  u'an  iintiT  I'ontainini;  a 
'.'ascinis  niixfurc  with  alioiit  10  por  cciif  of  CO..  Iiiiiiicdiatcly  after  fill- 
ing Ihp  liiniis,  ho  iiiaUrs  a  ray)id  forced  cxpirntioii  into  a  fiilie  provided 
\\ith  a  'al\c  ha\inu'  four  ojiciiiiiL's.  'I'liin  valvo  is  turned  throiiirli  a 
■nin|)le'p  circuit  diiriii);  tlic  expiiation.  so  that  four  fractions  of  the  px- 


|iiri'i 


1    lir  can   he  pollccted   in   nihher  hacs 


lecled    with    side    .iihcs 


o|iciiiii(r  opposite  the  foiir  oi)enini»s  in  the  val\e.  The  first  fraction  will 
rntitain  !i  little  less  than  10  per  cent  CO..  the  second  distinctly  less, 
Mhile  the  I'ourth  will  contain  the  same  as  tlie  third,  iiidicatini;  tliat  pipii- 
lihrium  lietwcen  the  CO.  of  tiio  alveolar  air  and  the  lilood  must  have  heeii 
.itfained.  Tiiis  (Isriii'c  tlieiefore  irives  us  the  Icnsion  of  CO_.  in  the  venous 
lilmid  of  the  lunns.  In  Henderson 's  method  the  rehreathiiiR  is  jier- 
I'liined  into  Kas  receivers  pontaiiiiiiir  <!  per  eent  COj. 
Tlii'sp  results  then   indicate  that   tlic  wliole   process  hy   which   C()j   is 


'icliaiiKed  in  the  Inn 


iff  use 


s  from  n  |)lace 


is  dejieiidi'iif  mi  the  law  of  j;as  diffusion;  the  (ids 
if  lower  to  a  phtce  of  hiirhpr  pressure,  and  does 


until  eiiuilihriuiii  is  attained. 


"4E 


I  it 


I  ill 


n  1 


ClIAI'TKU  XXXIX 
TIIK  roNTK'dl,  (i|.'  K'!.;si'll{.\TI(>\   ((■..nfdi 

THE  ESTIMATION  OF  ALVEOLAR  GASES 

r.v  K.  <i.  I'lAUd..  1!.A.,  .M.I). 

.M.'tliuds  Midi  as  tlijit  ..I'  lliildaiic  iiiid  riicstlcv ,  wliidi  ciilculatf  tli. 
iiiciin  pfrcciita(.'('  I'oinjxisitioii  oi'  tlic  alveolar  air  \>y  aiial.xsis  of  a  saiiipl. 
taki'ii  fn.in  U\v  end  nf  a  iHoldiiycd  fon-od  cxpiratidii,  <,'ivc  valiifs  wlii.l 
aiv  t.M.  liiirli  r.,|.  (■(>_,  jin.l  tun  low  for  (),.  There  are  several  reason- 
fnr  this:  (1^  III  the  time  taken  for  the  i)rolonir,.,i  ,l,.,.j,  expiration  ai 
appreeiaMe  ainoimi  of  CO.  \\ill  he  -.'iveii  olV  hy  the  l.leod  to  the  alveohii 
air,  and  oxvir.Mi  ^^ill  he  ai.sorhe.l  -  that  is.  the  sample  will  not  eontaii 
the  same  p.'iv,  iita-es  of  Co..  and  <  t,  at  different  stages  of  expiration 
(2)   The  portion  of  the  tidal  air  whieh   reaehes  the   alveoli   dilutes  th, 

alveolar  air  and  thus  eaiises  tin'  aiiioiint  of  CO,  j.'iven  otT  hy  the  hi j  t.- 

vary  diirin<r  the  ditTerent  jihases  of  resjiiration.  If  we  iiear  in  mind  that 
the  tensions  of  CO,  in  the  alveolar  air  and  in  the  I.I.mhI  leaviiif;  the  hum- 
are  always  th.'  same   (pa-.-  :U:!).  and  that  th Hire  fall   in   CO,  tensioi, 

iji  the  alveolar  air  occurs  <lurin<,'  inspiration,  then   it   is  clear  that  tin 

hlood   in  the  i)uli ary  capillaries  must  have  a  niaxiiimm   tension  a:!.l 

load  of  CO,  at  the  end  of  expiration,  and  a  miiiimnni  tension  and  load 
nf  C0_  at  the  end  of  inspiration.  Aecordinsjly.  the  averaire  of  the  pei- 
centau'e  of  C(  t,  ami  O,  at  the  end  of  inspiration  and  expiration,  as  d( 

teriiiiiie.l  ])y  the  llald: -rriestley  method  or  hy  any  of  its  modilications. 

mii'it  fail  to  'j'yc  the  correct  mean  tchsion  ,,f  these  !:ases  in  the  alveolai 
air  diirini.'  I'xpiiati.ni.  The  error  which  makes  the  CO,  hi<rher  than  i- 
should  he,  makes  the  pcrceiitaiic  of  ( t,  less  than  it  should  he.  These  in 
lluenees  taken  aloiiy:  with  the  fact,  which  will  he  shown  later,  that  tlh 

evolution  of  CO,  from  ihe  hi 1  is  iclatively  more  rapid  at  low  than  at 

hifrh  tension  of  CO,,  indicat.'s  that  the  hlo,.d  in  the  i>nlmonary  ea|>il 
laries  dm  ini;  inspiiati.m  must  contrihule  a  uneater  part  of  the  CO 
excreted  duriiiLr  a  respiratory  cycle  than  that  in  the  pulmonary  ea|)il 
laries  durin-j  expiration,  and  moreover  that  a  irreater  jKirt  of  the  Co 
excrete.l  must  he  cvolv.'d  at  a  tension  which  is  helow  the  mean  t.'iisioi 
of  the  CO,  presei.t  in  the  .•ntir..  time  of  the  expiration  We  eoihluth  , 
theref.ire,  ilial  the  avera-e  tension  of  CO,  in  the  alveolar  air.  <Ieterminc.i 

■Mi 


Tin:  I'oNTKoi.  (IK  Tin:  i<i:si'iuati(in 


.-{4.1 


li>    tlir   iii-tllal    li'iis'iMii    iiikIi'I-   wliirh   llic   '^ns   is  c 
liss  tliini  tlic  ;i\('r;iy:f  tciisiciii  (»f  CO.,  ill  till'  iilv( 


l\C(l    fliilil    lllc    IiIiiikI.    is 

liir  air  duriii)'  flic  tiiiii' 


it'  :i  ri'spii'iiloi  V  cyclf. 


In   tl 


IC    CilSC     (> 


r  (),  til. 


HKJitiiuis  iirc  (iilTcrciil.      While   llic   diliitiiiK 


ll'cci  (if  the  ahi'dlar  tidal  air  is  iiiai-i<c(l  in  altci 


the  aimiilMt  (if  ( 'O, 


-ivcii  (i(T  (luriiii.'  tlic  dilTciciit  idiascs  nf  a  rcsiiiration.  it  can  iiavc  little 
iiilliiciicc  (III  tlic  aiiKiiiiit  (if  ().  taken  up  liy  llie  lildod  iiiider  iKirnial  e(in- 
liiiiiiis.     Tliis  is  evident  fnini  a  stiidv  (if  llic  dissdcialidii  ciir\c  (if  li 


:;liiliil 


:  |iayc  :!s:!i.   which   sIkiws  that   at    tciisidii 


IXC    (l.'l     Mill 


eiiiD- 
I.    M''  the 


h.iiidyhiliiii  is  |iraeticall>  saturated  with  (>..  Since  the  tcnsidii  (if  (), 
ill  the  aUcdlar  air  under  iidriiia!  cdiidilidns  is  irrcater  than  fi.")  iiiiii. 
!tri-10ll  IIIIII.  i,  the  rate  (if  alisdrplidii  of  (>  must  he  piact  icall  y  iiiaxiiiiai 
'luriiiu'  the  respiratdi-y  cycle  tliat  is.  it  will  imt  chainrc  at  dilTereut 
iiliases  (if  it. 

While  the  rcjatidiiship  d|'  the  aKcdlar  'jases  is  ciint iniially  chain:in!.' 
.it  (lilTercnt  stages  (if  the  respiratury  cycle,  their  mean  rclatidiiship  fur 
pcridds  inclndiiiy  sc\cral  icspiratidiis  or  fdr  cdinplete  res]iirati(ins  is 
iiidic  (ir  less  c(iiislaiit,  licinji   cdiitrdiled   l)\    the  t.\  pe  nf  the  iiietalidlism. 

iiid  liiatheiiiatically  expresseil  liy  the  n  spiratur.x  (pldtiellt  (  pairc  "'47). 
Tlic  averau'c  relative  pcrccntaL'es  df  the  twd  lmscs  in  the  ahcdlar  air 
MHist  thcrefdre  he  the  same  as  in  the  tidal  air.  In  the  alvcdlar  air  c(il- 
i.cted    liy    the    llaldane    iiiethii(l.    hdwexcr.    the    almxc    faefdis    cause    the 

I  spiratdi-y   (pKitient    td  he  less  than  that    in   the  tidal  air. 

These  jxiiiits  liaxc  I n  insisted  upun   liecause  iiiiKdi  df  the  kiniw  led-rc 

Ml'  the  u'ascdiis  exchaime  I'ctxveeii   the  IiIikkI   and   the  air   in   the   liiiiu's,  as 
ell  as  the  ('(infrdl  (if  fcs|iirati(ili,  has  lieeil  liuilt   ti))(iii  data  (ihtaiiicd  liy 

lie  lialdane-I'riestly  iiicthdd.  and  in  cdiisideriiiy  this  wmk,  which  w c 
-liall    (id   in   suliseipieni    payes.    it    is  adxisahlc   that    we    lie   av\are   df  the 

iiiiilatidlis  (if  the  iiiethdd  eiiiphiyed.     The  liiethdd  has  lieeii  all   invahialile 

lie  f(ir  (ii)eiiiii'/  up  a  hitherld  entirely  uncNphired  lichl  df  research.  Imt 
||'iv\.  the  pidiieer  « (irk  ha\inu  liceii  ddiie.  we  must  cmphiy  mcthdds 
'   lii(di    will   enalilc   lis   tn   exphirc    iinire   e\actl.\. 

An  Accurate  Standard  Method  for  Normal  Subjects,  The  imist  aeen- 
ilc  liiethdd.  and  (iiic  free  fidiii  many  df  I  he  thedretic  eri'drs  ])reselit  in 

! Iliers,  depends  dii  the  rclatidiiship  fdiind  tn  exist  lictwcen  the  diltit- 

-  '•(^'••'■\  of  tile  air  in  the  dead  space  iscc  pauc  :;(rji   .|||,|  ih,,  knuwii  per- 
■  'iitaL'c  cdiiipdsilidn  (if  the  aKcular  air  in  expiralidiis  which  are  (if  var,\- 
'n;  depths  lint   <//'  iiimil  mul   unruKil  (hirdlinii   and   wiiieli    fdlhivv    iKirmal 
-pir.itidiis  (  \{.  <;.  I'earcc>. 

Ill  Ills  iiictliiiil  tlic  Milijct  is  iiiihIi,  1,1  l.ic'illic  llir,'u^li  \;ilvi,s.  wlii.li  :ii|f.,ni.,ti,.,,)|v 
I'lirntc  111,-  iiiv|,i,.,,,l  In, Ml  till'  i\|.ire.|  ;iir.  The  cs|,ii,  .1  iili  is  !,,,1  inl,,  -i  )„I„.  ,.,,ii- 
•'"•1   «itli   l\Mi   s|.;;iiiiir|ris   In    Uv  I   tliiivVMiy   s| ,,|wii,ks.      Tlic   s|.i iclcrs   ;dc   nf   (h.. 


»  9 


i  m 


l!!!l^ 


:!4(; 


Tin;    KI.SI'IHATION 


ll 


i,     W  r 


ii;i>l  Kinjrl,  iv|„.,  ,,„(•  l.riri;:  ,:,i,:il.|.'  cf  li..|,|iii-  t.Mi  litn-,  :iri.|  tlic  (itlu-r  urn-  aiul  a  liiili 

Tlir  exact  tiiiic  iliiiin-  wliicli  nii-  .iii.i-  i>  i. r,li,l  l,v  tli.'  small  spire, mrtcr  l.y  means  m1 

a  ;;rn„ve,l  dial  on  the  axis  ot  ll.e  li.|,  mi  ulii.l,  a  tinea. 1  works  over  a  system  of  piilUn^ 
ami  any   ■novcnient  is  ace.irat.Iy  i,  .-..Lira  l,y  a   wiitin^'  p,)int   on   tlie  snioke.l  paper  of  .1 

''"""■      ■'''"'   >|'ii'. meters  at Hie,-te,l   s,,   that    tlie   an    nnieiit    niiiy   I.e   ilireele.l   in   Hi- 

I'lne   I'ullouii,^   »;n.:      (1>    ll, ,,„:;;!,  c,,,-!,^    ;   ;,,  ,)   j  ,,ii|,i,l,,;    (o-,   ,ii,.,.,.,|v  tlinm),'h  l.oil 

""■'-•'  '"""  ''"■  l^"^''  ~1'' ler  l'>;    ilie  |.in|. -e  of  .oileelin;:  a  scries  of  expiration- 

an.l  ,::>  tlnon:;li  Cock  /  .Inctly  ii,i..  tli.'  -111:111  -|.irnniclci  tor  catcliini;  a  sin^'lc  oxpira 
''"II-  III  all  e\|icuiu.iils  tlic  liiM  Mlliii.;  ,  t  ill,.  -pii,,riiet,'r  is  rejertc.l.  so  that  the  (lea.! 
space  of  lie  -|.ii.  n,rtcis  is  nil,', I  uitli  air  of  a|,|.i,i\iniati'l>  ili,'  same  .•,.nip,,sition  n»  in 
111,'  siir,,.,',lii,^  , Ai'ii.ni  'IIS.  T!„'  lini,'  i-  ni.nk.'.l  In  M',-,in.Is  l.y  a  tim.'  ■l,.ck.  The  rcspira 
l,.iy  iii,.\,'in.  lit-  ai.'  1  .■.■., r.l.'. I  l.y  a  |  1  .•iini..L:ra|.li,      ,  I'i:;.  IJs.) 

'I'hc  sii!.j..-l    is  I'l.iii^lii    into   1,  s|.iialoi,\    ,  .|inlil,i  iiim   l.y   liaviii;:  him   breathe  throii-li 

111.'    \aK,'s    f,.r   a    |'.'ii...l    ..f    tin,.'    I',.f,'i,     tl liS,'M  af  ion.      'Hie    r.'spiratory    niovemonfM 

,liirin.;  this  tinii'  ;ir,'   ie,-,,i,|i.l   wliil,.  :1 -ksari'  in    l'i.-,tiori    7.     When   the  ol.servati.m 

is  staite.'.  III.'  .-o.-ks  a-.'  tmne.l  int.,   l'..siti..ii  J'  .Imiiii;  tli,'  lime  an  Inspiration  is  hoin^ 


1 


.'■>  ^ 


'TV" 


:rM^r^^ 


\/'/ 


^^^m::^' 


1: 


l-'iK       1-'*  Alt.!!! 


If!  I 


i:t    ■!    i':(itr>    aii.i    >  .'r,n. .  ti-i;';    of    1'.  .trot's    int  i!:-..!    fni     iiu-.T-.ir«  incut    ni    (i* 
"f   alvc-I.ir    ;i!r    iii    Por;\:.iI    ■JiiI'Mt;-. 


111...!.-,   s..    iliat    til.'  cs|.iiati.ii.s    ulii.l,    f,,|l.,u    may    I lle,-to,l    in   the    laigo   spiromct,  r 

After  al.'.iit  t,'ii  i,-|',,al|.iiis  ,a  ."ii.i.'.l  nmiili.'r!  have  1','cn  .w.llert..,!,  the  cocks  ai' 
tiniM'.l  1  '  I'.sitiiiii  ,;  .liniii-  an  in-i  i'all.iii.  aicl  a  siin;!..  .h.'p  expiration  is  colh^.'t,  ' 
m  th,'  small  -|.ir,in,.t,'r.     Ii,  ,,i,|,.r  that  Hi,'  thn,'  of  this  niav  l„.  ihe  sann-  as  the  norm:,' 

•  xpiraln.n.   it    is  n,.,essai\    t..  ,|iii,k.'n    It.     This   i-  re  or  less  a  eham,'  pro.'piliire.  Lu- 

with  a  littl,.  tnilnhi-  -h.'  ,.iHrai,,r  ,  .-ni  ,  !,.s.'  1 1,.  -i,,p,',„.k  »ilh  snlli.-i,.,,!  ai'Ciiracy  i' 
iiiterrnpt    th,     <1,  .p    ,'\pirai|,,n    .ai    •]„■    .  n.l    ,.t    tlo-    n,,rnial    expiratory    time.      Slionl 

tliir..  I.,'  .niy   yr..ss  variation   fi  m   111..  mal  .■\piralory  time,  th,'  sanipl..  mnst   lie  .-..I 

I.-.I..1    a-ain.      \..i    in fi, .10.111 1\    ih.'    i.i>piiati..n    iniin.'.liately    pieee.lin^'    the   i.xpiratl.c. 

Into  ih.'  -mall  -|.ii,iiiiit,'r  is  \ari..l   iii\"liii,l;ii  ily  l,y  th,'  snl,.jei-t  on  ai nut   of  his  li.in. 

a»ai,.    lliai    th.'    f..ll..»  in-   I'xpliatl'.a    has   t..    1..'   .I,,.,  ,',„.,|    an.l    finioken,"l  :    this   .-an    ! 

partially  ,i\,'i.  .0.1.'  l'\    ;:i\iiiu-  him  th.'  .-i^nal  to  l.reat! iit   .Icply  after  he  has  actiiall- 

I'cuun  I..  .'Vpii.'. 

"cteriniiiali.il.-  ai.'    ina.l.    ..f   tl,..   av.ia^e   m.Iuiii.'   .,f   th,.   ii,hil   aii     ,,.,■.   air   in   larc' 

spir,.m.l.'r  .ihi.L.l  l.>    1 1..  r  ..f  l.r,alll-^.  ,.f  if!.'  i,,l,|,,,,.  ,.,.l!,.,.i,',!  I'mm  th,-  .i.ci'.  .  ■. 

piration.  aiol  of  ilo-  p.-rcnlaL"'  .  ..ii!i...-ll  i.m  <.f  th.-  ti.lal  air  an.l  that  of  tli.'  .1.'.'; 
.  v|.irai|..i!.     A  .  lit.  1  i..;,  f..r  .l.'i.'iiniiiliiu'  .» lutlor  or  not  ilo'  pro.'e.liir.'  has  l.een  carri.  : 


Tni:  (oNTKoi,  OK  THK  ri:si'iration 


347 


,t  .•  ir.ctl}-   is  till'  irs|.ir;itiir_v  (|uolitri(    (latin  "f  CO,  rxcieti'il  to  O,  alJKnrlicil).     Knr 
•  :i-."ns  wliic'li   arc  set    I'nrtli   hIkivp.   tin-  i|Ui)tii'iils  slmulil   lip  .•i|i|iriixiiii:iti'ly  ('i|iial   in   the 

.,  !■  idlli'i'tril  ill  llii'   laici'  aii'l  in  tlir  small  >|.ir. ti-i>;   it'  liny  a  I'l'  nut   sn,  tlic  ccjiiili 

■i"M»  lit'  ttii'  iiic'IIiimI   litiM'   iKif   l.(rii  roiiiitly  raiiii''!   nut. 

Siiicr  tlie  (trail   space  ami  tlio  tir(riiiji    i-(ini|r(isilii,ii   of   tlic   ahoiiliir  air  iimlor   tlicse 

ii.litiiiiis  may  lie  i-misiilcri'il  ciiiistaiit,  tin-  |irii<'iilaj;i'  cnmiiiisltinn  iif  tlii"  deep  cx[iira- 

Ipiii  will  (lilTor  fiinii  tliat  of  tlio  iiiixoil  saiii|il("  iif  scM'ial  iinrnial  rxpiratimis   in   ppiipnr 

li.iii   iis  tlie  ilcail   spair  exerts  a  jjieater   itiliitiii;;  etTeet   in   tlie   small   tlian    in   tlie   laiye 

\|iiialiMii.      Tliis   li.'iiii;    till'   ease,   the   ilatu   I'litiiine.l    ran   lie   eiiinliineil   aleeliraieally   to 

_i\e    eitlier    tlie    eapaiity    nf    the    air    passapes    iir    tlie    |ieieenta;;e    eumpiisitinn    of    the 

l\r.ilar  air. 

I.i'l    A   r_-  ameunt  nf  aii    in  larye  expiration    (small  siiirnmelprl, 

Ai  -z  amiiiint  ot'  aii    in  small  or  iiiirmal  expiration   (tiiliil  air"), 
H     -.:  tlie  perei-nta^e  of  CO    or  (),  in  tlie  expireil  air  of  l.'irjje  expir.-itinn, 
l!i   ;=  tlie  perc'enta;;!'   ''f  ('0._  or  (V   in   tlie  expirP'l   iiir  of  small  ex|iiration, 
X  -:  tlie  rapai  ity  oi'  tin    ilenil  space, 

y  rrt:  tlie  aveia^e   priiehta^'r'  of  ('(!_  or  O^  ill  til'  alveolar  air;   then. 
\  •■  11  =  (A  -  x\\  iiii.l  Ai   ■:  I'.i  _-  CAi    -  x1y. 
s..l\iii^'  tliis    for  \.   y   lemaiiiin^   loiistaiit    iimler   tiie   same   pnysiolo;;ie   eomlitions,   \m' 

\  ■  Ai  ■  I  r.  Hi  1     ,      ,     , 

^-.   tlie  ilea.l   sp; 


A  X  n  -  Ai  •<  Hi 
AxR- Ai<r! 


Or  soh  'M^   fill    V,  «e  lia\( 


J ,   tlie   mean    prneiitaue   of   CO.    in    tile   aI\eo|ar   air.      fn    ea«o   tlie 


A-Ai 

■I  ;mI  Mpaee  for  O,  is  ile^ireil.  1'.  an-l   iii  must  lio  maile  to  eipial  the  O,  alisorlieil. 

Clinical  Method.  Tlie  use  of  tlie  UynmiriJiiili  aiul  inicuiniisri'apli,  ami 
I'liei'  eoiii))lieiiliiiir  fjtctors,  iiiiiki'  tlic  iiictliml  lis  jiist  (lescfilu'd. ruiito  illl- 
|i|,ietie;ibl('  fiif  cliiiieal  iiiiieodiiri".  liut  till'  Use  of  tlio  same  appiifatiis 
■' Ith  llic  follow  iiiu  iiiiidilieiitioti  will  yield  satisfaetot y  losults  for  most 
■liiiieal  pufjioses.  The  iiatietit  is  iiinde  1o  I'esjiiic  tln'oiierli  tlio  valves  fof 
,i  shoif  tiiiip,  aftof  wliieli  (lie  olisorxi'f  folloefs  ji  siti<ilo  expifatioii  in  ti 
-mall  s))ironiotei'  liy  tiifiiiiiir  tlic  stopeoek  ffoiii  Position  1  to  2.  A  saiii- 
ple  ><(  lliis  is  taktMi  for  analysis,  and  the  spii-oiiictcf  is  asaiii  eiii|)tiod 
lid  a  sei'ies  of  siieeossivc  saiiiiiles  of  deeper  expirations  takoii.  This  is 
'li'iie  hy  dircetiiii.'  the  |)atieiit,  after  ho  h;is  st;ir1ed  to  lircatho  normally 
into  tho  spiromotor,  to  hroatho  more  deeply.  The  aiiiotnit  of  air  eol- 
!' efed  in  each  oxi>iration  is  coiitfolled  li,\  the  olisorver  liy  elosiiii;  tho 
-'iipeoek  when  tho  ilosirod  vohmio  is  olitaineil.  I'y  tliis  means  ono  oan 
'iillect  several  oxiiirations  ditVerino-  from  ono  another  Iiy  iiioroasiiii.' 
■  iiMnlints  lint  all  oeeiipyiiio;  the  same  time.  The  samples  of  the  various 
'xpiratioiis  are  eolloeted  in  a  series  oi'  luiinlierod  saniiilintr  syrinoos,  and 
'ie  oaseons  oomitnsitioii  of  eaeh  is  deleiiiiinod.  When  the  poroenta<;e 
I  i  <'0^,  or  O.  in  oaeh  expiration  is  |ilotto(l  on  cross  section  ])aiier  on  the 

dinates.  with  th.o  volume  of  tho  expirations  in  ctr  mi  the  ,n!)seiss;)>^  a 
i  \  perholie  eurvi'  should  li(>  ohtained.  Atiy  marked  deviation  from  such  a 
iiirve  indientes  that  some  error  has  hceii  made  in  takinf^  a  sample,  and 


5l 


I  :  liiiii: 
i  -i    11, 
m 


■f  ! 


\\ 


m 


m 


ill 


\  III 


I     i 


.i      II 


'  i      ■'■t 


Ml 


H; 


348 


Tin:  Ki;si'ii(\Ti()\ 


llii>   nlisciviition    slmuld    lie    (liM-iir.lf.l.      Tlu'    .lillViciit    .il.s.-rvatii.iis    a 

"!'■"   ' '''"'■'!    i"    ""•    t'iniiiihi    uiv.n   uii    i„i;rr   :!4;,     Tlir   .l.'t.Tiiiiiiiiti^ 

ol' the  (•(»,  pci-ciitji-f  .i|- cxiiiifil  iiii'  i>  Ml  siiiipl,.  tliiit  ;i  iiuinlu-r  (if  spc 
ni.'i.s  „(  \,n\ii,u  .Irptlis  (.I'.'x|.ii;,ii,,ii  .•iiii  !„■  liik.'ii  iiiiil  iluis  nmiiy  jMiiir 

""  'I '"^''  il<'t<'riiiiiu'(|.      K,,i-  Ihf  nidsl  ;ic.-uiiit<'  rcsiills  it   is  in  ^(.i.'r,: 

''•'^'   '"  '•"'iil'ii'' 1>    tli,,M.  ...Npiiiitioiis  u|,i,.li   .lilVcr   tVuiii   ..ML-  aiKitli. 

Iiy  at  Icasi  (i.:;  pri'  ,-,.nl  in  (  ( t ,  ami  l.v  al  least  'JdO  <•..•.  in  voliiiiii'.  Th 
il''p.'ii(ls  „u  Ilic  lart  thai  tiic  .lilntiiiir  cllVct  ,.r  tli,.  d.'acl  space  in  rniip 
iiiu  tlif  pcivi'iita'^r  i<(  CO  ill  111,.  ,.xi)ii(.(l  ail-  tioiii  that  in  tlic  alvcul,, 
ail-  is  u'l-.-atiT  in  n-lativ.-ly  Miiall  r\|,ii-atiniis.  If  nioi-.-  exact  work  is  <\. 
•<irn|.  iIm-  (»,  (-.Hit. -lit  ran  he  (Ictciiniiic.l  on  eacli  sp.-i-imiMi,  tlic  rcspiratni 
'lilniicnt  cali'iilatcl.  ami  ,,nly  tlnis.'  .-xpiiatioiis  w  liid,  sln.w  fh,-  sain 
irspiiatipi-y  ipictii-iit  (•(iiiiliincd. 

Ill  tlic  taMc  ca.-li  oJiMi-valinii   is  i- parrel   uitli  cadi  ni'  the  otiicis  i 

all   imssililc  cdiiiliiiiarnuis. 


Pi 


1-1 


\'i.   (pK 

IIIIM 

K-^i.in  '. 

1  \i-li:i 

ION 

All: 

1 

l.'.ii 

U 

<:.".7 

7."in 

llL" 

l> 

Ilia 

n  i:  1  I  \T 

I  Cl      IN 

!  \nia,i> 
mi: 

:;.Iii 
.■;.(;i; 
i.iiii 

l.-.N 

i.:;ii 
t.lii 


\.W  I  I  1711 

■''■•"  :  "1^  ."..J7  |v.i  Is:;  -11 

■'■H  :  "i.I'i  I.HL'  :  nil  I      11(1  |s| 

."..!<;  I  \:>S  I      t.H-  I  iri  i:;7  :     171 


<iiiii-i;il   :iM'i;ii;r    r..i    CO..   in  ;il\|...l;ii-  nil.  ."i.l::. 

'-mml   :,,,.,:,:;,.    I„,    ,|,.;„l   .|,:„-,-.    17l'.      I),-:..!   >,,:„,■   in    M.hrs   in   tl.i.  cn,.,- ml    u 


■:-fli 


Anntlici-  iiK'ilioil  «|ii,.||  l,;,s  I,,.,. II  siioycstc.!  for  clinical  piii-ixiscs  i 
tlnil  ,i|-  JMi-sch;  tliis  ,•,,||sist^  in  liaviny:  tlic  suli.jcct  ln-caflic  several  linn 
ill   iiii.l  .Kit   .if  a   small    lia-.      It    is  assiim,.,!   that   after  siich   i-esi)irati.. 

'' i'i|>"^>iti if  th,-  air  in  Hi.-  lia-r  will  h, me  llie  same  as  tliat  in  tl 

ah.-.ili.       Alth.iu-li    this  -,,  ,|„„|,t    till,',    it    has   li.>eii  sh.wn   that    tl 

Miclh.i.l   is   fallaci.iiis.   1  til,.   (■(,.   tension   .Ictcrmiiied   in   this  ^^;^ 

is  mil  that  .if  th,'  ai-t.-rial  lilo.i.j  al.ui,'.  Imt  is  the  avera-.'  hetwcen  it  ai; 
thai   .if  the  veil. Ills  lilo.i.l. 


S  3i 


?|: 


(•IIAI'TKi;  XI, 
TIIK   ((tXTI.Mil.   (H"    l.'i:s|'||{.\'|'|(t\    .C.ntM 

THE  NATURE  OF  THE  RESPIRATORY  HORMONE 

Tilt'  piiM'ticiil  iiii|>iiif;iricc  ol'  tlic  iilisci\  jiticuis  ilcsciihcd  in  ihc  furcirdiiin 
rli:II)t.MS     ill     till'     ilivcstiv'.-itioli     ol'    till'     l-.'l;ifii.iislii])     lictwci'll     ( ',,     (.f    tllr 

I  I'Miil  and  it'spiiiilipiy  iictivi1\  will  now  he  |)laiii,  jiinl  it  ic-iiaiiis  tor  ii> 
1-  (•(insider  tlic  pliysiidooi,.  ,.\  idcm-c  that  Midi  a  r('lati(pii>irip  exists.  In 
rhi-  lii'sf  ])lac(',  let  IK  ((Misidcr  the  behavior  of  the  aciJ-base  equilibrium 
during  conditions  of  abnormal  breathing — hyperpnea  and  dyspnea/ 

As  CO,  accnninlatfs  and  O.  Iiccunio  used  up  in  a  ('(infilled  space,  the 
'  'iitliinu'  liocdincs  iiiteiisilied.  In  scaicliiiiL'  I'di'  tlie  exact  cauM'  (if  tliis 
■■^Vcct,  we  iiinst  first  dl' all  ascertain  \\lietlier  llie  |i>  perpnea  is  ,\\u-  u>  tlie 

■  !■  Ilcinicy  of  ().  (ir  t(i  the  acciiiiiiilatidii   dt'  ('•>..       Many   dt'  the  e\|>eri- 

'  Ills  lieariiiL'  (111  these  pi-dlilciiis  can  lie  iikmc  >atisractdril.\    pert'eniied  (in 

'M.iii  than  (111  l.ilioratdry  aninials.  hccaiisc  anesthesia  is  ikiI   necessary  and 

!m'   siilijecti\  (•    syiiiptdiiK    e\])eiiencc(l   are    (if   ureat    value    in    tlie    intiM'- 

irctatiiin  of  the  results.      If  an   individual    is  placed  in   a   larjio  fiir-tif.di* 

'.iiiilier  cjOOO  liters'  ca|)acity '.  and  the  depth  and  rate  df  lireathiiiL'  dh- 

■  rvi'd  as  the  CO,  acciiinnlafes  and  the  ( >.  Iiecduies  used  up  in  the  air  (if 

■'<■  chanilier.  im  distinct.  chaiiL'C  in  respiratiiin  will  lie  dliserxed  until  the 

I  '»,  pcrcentaife  (if  the  air  has  risen  tci  alnidst   :!.     AIkixc  this  jidiiif.  how - 

er.  the  hyperpnea   lieennies  iikhc  and   nioic   i)r(iii(iuiieed,   until   finall.v. 

'icii  the  CO.  ix'tceiifa^'e  has  risen  Id  alidut  li  and  the  O.  )>('rcentaL'e  has 

'  'Hen  trt  13.").  it  li iiies  unliearalile  i  dyspnea  V     l-'nnii  the  results  nf  the 

"■i.niinjj  olise  -v  atidii  ahiiie  wc  cduld  nut.  lid\V('\cr.  decide  whether  the 
'    •itatidii  (if  the  respiratdi\    center  is  due  to  the  deficiency  df  O.  (ir  td 

■  '  increase  df  CO..  If  the  experiiiicnt  is  rc|)cate(l  with  the  ditVerellce 
it  the  CO,  as  it  accuiiinlates  is  alisurlicd  li.\  sdda  lime,  iKi  liypei'i)iiea 
M  (Icvchip  e\('n  when  the  O,  is  as  lnw  as  in  the  |ire\i(ius  experiment 

''■'■  may  eoiielnde,  therefdre.  that  in  the  first  i\|ieriiiieiit  CO.  iirrKiiiiiInlioii 
'v/  liarr   artul  ii.i  tlif  ri  siiinihirii  stiniiilK^. 

rill'  same  cdnclnsidn  is  arrived  at  as  a  result  df  dliservatiiins  nn  indi- 
lual.s  Paused  to  lircathc  in  a  ninre  cdiilined  space  as  intn  a  ruhlier  ha'; 
ahtiilt   2'2ri    liters'    ca|iacit.\.      I'lidcr    tlicse    cdnditidiis    li\  )iirpiica    dc- 

■■' ■/■■■'7'".<i    iiii-.iii-    -IJuhllv    iiurr.i..,l    l,i,-.iilirit;    ,/>,,/■,;, ■,!     l:,l.,.r,,i    l.r.ai hiiiL'.    l.-n       .1    mmIi    .;v, 
■    I.  iilil;,li..n    I,,    ni:iiiit.ii.i    lit,;    ,i.,^;.,';,j      111,     1, -:,!•.     ,|     , ,. -iilVi,  i,nl    Li .   il  lii'ii;. 

34!> 


>-\  i^'3! 


i!.l:ii!i 


I  irfji 
i  11  i.  i 
i  I  ill  m 


;!.■•() 


'IMI      HI  >ril(\  IIUN 


I        ! 


i  I' 


r 


iJ^ 


I 


i  >■ 


i    I      ■  I. 


vcldps  iiioif  i;t])'ui\\  iliaii  ill  lli,'  l.irjiv  ciiliiii.'t,  iiiid  a  IukIkt  iMi-.Miila.j. 
'  It)  ])fT  .-.•lit  1  .if  CO.  can  lie  l-.l.-ialcl.  Tlml  in  this  caso  also  .Icli.'icn.-y  -^ 
<>,  is  nut  n'sponsiiilo  l'..r  'li.'  Ii\  |.cr|in<'a  .'an  lie  shown  l.y  ri'iM-tilion  of  \\r 
'■xprniiicnt  oitli.T  with  an  ..sc.ss  of  (»  in  thr  I,a^'  or  with  al>sor|)tion  of  tl  ■ 
i(»,  liy  scla  JiiMc.  Ill  the  fornnr  case  li,\  pel  imca  will  cl.'\f!n|)  as  iisii,i 
\\liil('  in  the  Jattci'  it  will  not  Mi|ici-vcnc  inilil  the  ))i'iccntav'('  of  (),  h,; 
fallen   h.-hiw     |(l,  « hen   eyaiiosis  liri'ciiies  inarUe.l.      In    tact,   seme   peoiil, 

lie.Minio  cyaiic.Nc.l  ami   in nscii.ns.  an. I  .-ollaps,'  imuI.t  tli.'s.'  .•.m.litioiis 

I'.'forc  thcic  is  any  lopiiat.oy  .listnrl.aii.-.'.     A   p.M-uliaiity  ..f  the  cIT.t- 
"f  (»,  ilelicicn.-y    is  that    the   person    nia\    lie   iinauaiv   ..f  tli.'  seriollsiiev. 
nf  his  c.Mi.lili.in  :  in.lee.l  he  may  1..'  s.mm.'u  hat  stimulate,!.     The  .■..n.-liisi..i 
may    he   ilrawn    that    <h  firli  iini   t,(  ().  p,  ,■  s,    ,  ,ni   .«,•,.    ,is   „    n.<i>ii;il'fr 
stimulus  iinhl   wh,  ii    It    Is  so   i.rlninr   as   tn   ,;iiis,    ollnr  s.rlo".;  siitnplo,,,.- 

This  coni-lnsiiMi  .lues  not  in!.'  ..nt  an  imp.olaiit  inllu.'i •■  (».  (lefieicn.- 

in   iin-reasiiii:  tli.'  e\citaliilit>    ..f  lh. nt.T   t..\\ai-,ls  CO..      I'n.ler  .iiil^ 

ii.ir.v    e.,ri,|iti,.ns.    Iiuue\,r,  t),,.  .-ciilir'  is   tar   iikmv  s,nsiti\e   towards  -Hl;!' 
ehanei's  in  th."  CO^  i).>rcentaire. 

Th.-r.'  IS  an  .ilivi.ms  r.^asun   why   th.'  a.ljiistm.'nt    uf  pulnmni.-  ventihi 

ti.ni  shonl.l  not  .h'p.'n.l  ti| haniri's  in  O,  supply  to  th,'  respiratory  <•.■! 

t.-r.     If  it  \ver.'  s.i,  many  .ither  tissii,'  activity's  ami  oth.'r  ne"\i nl.T- 

N'.ml.i  sutler  fr.im  lh.'  O,  <|e(i.-i.'iicy  li.'f.ir.'  th.'i-.'  was  time  f.ir  the  hreail 

in^'  t.i  Ii.i-.JMI.'  stininlal.'.l  siiftici.'iitly  t ak.'  ^...i.l  the  l.iss  .if  O,.     A-  , 

iiiatl.-r  .if  fa.-t.  h.'a.la.'h.'.  .Ii//in.'ss,  naiis.'a  ami  even  faintiliir  iire  alum- 
e.-rtain  to  lie  .■anse.j  \vh.'m'\..r  an.\  miis.'iilar  .'x.'i-.-is.'  is  attempt.'. 1  in  a 
alm.isph.'r.'  .-..ntaininu  :\  .l.'ti.-i.'ii.'y  .if  (^  Imt  no  e.^.'css  of  Co.,  i.-i".  nn.ii: 
tain  si.-kncss  .  An  a.le.piat.'  O.  supply  .if  the  h.i.ly  is,  tli.'i-ef..re.  iiisnr.  ■ 
li.\    chaim-es  in  CO     t.'iisi.m  .if  the  iilo.,.!. 

Quantitative  Relationship  between  CO,  of  Inspired  Air  and  Pulmonary 
Ventilation.     Th.'se  r.-.nlts  sny-.'st.  as  th.'  n,'\t  stej)  in  th.'  investi^athi' 
«(  our  pi.ilil.'iii,  th.'  .!  i.riiiinati.in  of  th.'   .plant  italiv.'   relati.mship   h, 
Uw'on   th.'  CO.   per.'.'nta-.'  .if  th.'   r.spir.'.l   air  ami   the   amount    .,f  al 
lir.'ath.'.l     pnlm.iiii.'  v.ntilatinn   .*     That  there  is  sii.'h  a  relationship  Ii;, 

heeil   m.isl    Mici'ssfiilly   .I.'m strat.'.l   liy    |{.    \V.   S.'.itt,   who  u.scd  f..r  hi 

piirpiis,'  ,|,'.-er.'lir;it.'  .-ats.*      Th.'  tra.-h.-a   was  in.'cle.l.  throiiyrh  a   T-tni. 

provi.ii'.l  uith  valv.'s,  with  tiiliin-  l.'a.lin-  p,  a  lar-.'  h.ittle  and  a  (iad-Kr.r.'i 
spir.miet.r,  s.i  th„l  th.'  animal  lir.'ath.'.l  out  of  th.'  Imtll.'  into  tl, 
-pir.imeter.     tli.'s,'     tuu     \,r\wj:     al.Mi     .■.iiin.M'i.'d      Pi^n'tli.'r.       The     spinm 


,-       '         ''~''i"li"ii    Ih    -.llHulitu-    .Ir.-i^n    l„'tw.-.ii    i.nln.nni,     \ ,  ril  il.il  i.-ii    .-i.i.l    .ili.c.l.r    v.nlil.ii    ..      t 
I"^i'>i...l    l...t|.;,-,.    |,^,i„„,n-,-    ;,,ml.iin,    il  ",h,'\^,(,"r    valll.  ■|!r>,'iv,."''    '    '    """■■    '"    "'"" '"        ' 


M>.  I  .  rc  iiralc  anini.ils    nm-t    I,c-    i|.,,l    in    ih, -.■   i->|„i 
<  tivit.    ..f   ilic    r<  v],i,.,tnry   tent,  r 


|i.  11KMril«,    vin,.     .111i-»tllili.  -    111.11  k.-.llv    .1,1,1, 


«ii^iU:i 


Till:   i(i\lliiii.  Ill'    III!     Ill  vi'iii  \'i 


ill'  was  liiiiili'  III  rcciii'il   its  iiiii\  I'liiiiils  on  a  ilnmi 


•Ml  iliiil   nil  aiM-iirati 


iiiH'iii'N   111  till'  ri'>|iii'atiiiiis  w  as 


iMlll'll.      Si 


nil 


niMii'd  ol'  tiic  (Iri)tli  anil  tVi 
liles  of  iiir  wvvv  riiiiii\  cil  irum  ijir  liuttli'  liy  i;iiiuii(l-i:iiiss  |)liiiitr,.i.  syringes 
it  frofnuMit  iiitiTvaU  ilillilii.'  tlif  linn'  tlial  tin'  aiiitnal  was  i'i's|)iriii^  iiifu 
■  111'  liiliiiiL'. 


!  '• 


' 11 1  1 : 

'■-"[ 

,    ,    .    .    . 

T 

-4-4- 

i     1 

J                                            '  ~~' 

j 

i 

■■ 

I 

JM  — - 

1 

-I-  - 

i   :    ]        . 

'           \ 

f    '  ;  •  :  i  II  1  1  1  1  M  1  1  1  1  I  T-m 

_i- 

■^  "^  ;    ■  :  ■      ■■]■" ■ 

■      1    i  ,    ' 

,  j 

' 

i    !      '          -^ -  - 

400  Ip- 

\    ,         1 

§ 

J  ^ 

- 

■  --i 

1   1       ; 

r 

y 

^-i-. 

ZL 

r 

' 

^ 

. ' 

1  '  I  ' 

A                                        .  -j—          .   .      , 

'             i                                                           t           " 

j    i       '  •         \    /                     

t 

1                         .            jr 

,  :       J                   _L    "■' ' 

2  _ 

J             ■     -  '-- 

* 

:     !     1  ■ 

,/     X        ~ 

— u_ 

ti                           ~ 

uX-  -^        ^^     4 

100-3- 

'  ■     # 

1     '  i/ 

-^- 

.  Jr 

'     '     Mt 

S 

•M 

[~ 

i       Jt 

i 

f 

\m 

/OC    . 

^ 

'  1      '  jj--  •■  ■-     - 

JT 

_ ":     +      1 

. 

/\        ;  ;          II             -  1 

1 , 

-  ^^ 

1  ,     :    ,    '    '     '                    i 

.y 

1 

^^^ 

t' 

JM       IM»  ACIk         fl  II 

_^5 

i    r 

,  ,    ,    .^^.       .1 

i^__™e  H|"|     -n  B  _j._^ 

1 

z 

3 

^ 

5             fc 

I    » 


'■i«.     iJ'i.      l.'-ini[M.sii,-    (•■■ii- 

.il'iry    ics]..iT!-c    lu   i  I  1 .    II 

nnlin;it.  <        Ilu    prr. .  i 


\c  tK.t;iii;<il  iti.M)  liic  il.ua  ..II  -iMiMi  I  \in  ■  nil- III",  ^hnuing  iLt-  re 
1  I'll'  il.'i-.  r.  1  i.itv  ,Mt.  .\l..,  i-ru'  |..  i.int.ii;,.  c.f  CI).  |i,  tin-  iiis|.ir.-.I 
tile   Hiita-i'   III.-   II  111    an    pr     .mniti'       (l-'i..iii    K     W.    St-.ttt.) 


liL'  rt'siiHs  ari'  ^rivi'ii  in  tii' 


iic'i'iimpaiix  iiii:  iMirvo  (  Fi^'.  12!M,  wliifli  shows 


It  til' 


'I'o  is  a  |H'i'lri't  i-iii'i'fN|iiiiii|i'n 


if  the  bottli'  and   tlii'   |Hiliiiiiiiary   xi'iitilati 


lii'twri'ii  till'  < '( t  iX'i'i'cntnKi'  in  tlii' 


>ii.     .M 


ori'llNiT,     \\  lll'll     till' 


ittl. 


fillt'il 


witJ!  (>.   insti 


if  air 


-tai't    with,   fl 


If    ^!llih'     rrSlilt' 


"II'  (il)taiii('il.  sliowini;  tliat  tlic  ('()._.  i iiiimlatinn  almio  was  i'os|)iiiisihli' 

I'  tlio  liyiK'i'piii'a.     In  tlii'sc  casi's  tlu'  piMfi'iitaKc  of  O.  foniaiiiiiifj  in  Iln' 


111       } 


Il'r 


iJiil' 


:i:.L> 


■nil    HI  Ni'iii\rin\ 


-<v  sli'iii  iil'li'i   li.v  |)i'i  |iMi';i  liiiil  licroiiic  i'\l  itnii',  w  MS  I'm-  iilicivc  lliiit  ;it  «  liirl 
ililcct    cM-iljitiiiii  III'  till'   criitci'   1111111    <•.   ilrlirirni'V    is   pnssildc. 

I'lNpi'iiiiii'iits  111'  ii  siiiiiliir  l>|H'  liiiil  prc\  iiiiisly  lircii  iicrriiriiii'il  li.\    I'm 
tiT  iiinl  Ills  pii|iils,-'  Init   tlii'ir  ulijci't   wiis  imt  so  iiuirli  In  slniu    the  tIhn 
piir.illi'lisiii  lictH  cell  llic  CO,  ciiiiti'iit  111'  till'  ri'spiri'il  ,iir  iiiiil  the  piiliiinni' 
\  rlltilillinll   iis   III  ilcliiiuistriili'   llii'   I'luiMirCs   pinilnccil    ill    till'   sensitivity    •.: 
till'   ii'spiintciiy  ci'iiti'i-  ill  piii'iMiiiiiii;i. 

Possibility  that  CO    Specifirallj'  Stimulates  Center.     Al'ti  r  sliowiii' 

lliJit  ( '( >,  acts  iis  i\]]  I'Xi'itiilit  111'  liic  icspiiiitoiy  cclili'l',  tlic  (pU'stimi  iil'isi- 
«  lii'tliiM'  «(•  ill!'  jiislitii'il  111  the  assii  nipt  inn  tliat  lias  Imm'm  inaili'  ti'iitatix  i'l\ 

that   till'  artinll  ilcpi-llils  mi   flii'  raisiliLr  nt'  tin'  ('n   nl'  tllC  Mnnd.  nr  wlictllri 

it  may  ln'  a  spci-itii'  actimi  »(  tiic  IICO  animi  itsi'll".  Afaiiy  attempts  lia\- 
lieeii  iiiaile  to  ileciile  this  ipicstinii  e\i)eiinieiitail\',  llie  tfeneral  priiiei]ilr 
nt"  tlie  <'\|)crinieiits  heiiii;  tn  (Icti'imiiie  wliether  ( '„  nf  tlie  Mood  luii- 
l>aialli'l  with  the  CO.  eniitciit  of  the  i'es])irc(l  air  and  with  the  liyperpiie.i 
I'siiiir  the  ;iasehain  iiiethnd  (  paire  ill).  Ila.s.sell(ah'li  and  Liindsiraaril 
I'niind  that  the  hypeipiiea  prndiieed  in  raliliits  liy  lii'<'atiiin!;  in  CO^-iicl: 
air  runs  api)rn\iniately  parallel  with  the  increase  in  the  ('„  of  the  Idond. 
Iiiit  nil  account  nl'  the  experiiiieiital  ditlieiilties  enenuntefed  tliey  eonld  iini 
decide  wliethei'  chansics  in  C,,  are  nhiiu  i'espolisil>l(>  for  tiie  pfTccf.  TIicm 
a.ilhnrs  had  ini'vinnsly  deiiionst rated  tliat  chaiifjres  in  \\,  can  Iio  indiicii! 
ill  hliiod  leiiioM'd  fniiii  the  Imdy  liy  alterations  in  the  CO.  tension  witliiii 
the  ])liysiolo(;ic  limits.  An  increase  of  one  millimeter  in  V0„  tensini 
was  found  tn  cause  an  iiivrcase  in  ('h  nf  OIKHi")  ■  1(1'  (see  ])a}ro  27  i. 

1{.  W.  Scott's  experiments,  aliove  referred  to,  liave,  however,  yieldcii 
more  definite  results.  T.y  usiiii;  the  colorimetrie  method  for  determiniii'j 
C,i  nf  the  IiIihmI  (see  pa-re  •>"J).  it  cnuhl  Im'  readily  shown,  as  is  cviddi' 
from  the  talile  (Col.  S  in  talilei.  that  a  marked  rise  in  ('n  lieeiime  evideiii 
\\lieii  the  inspired  air  cniitained  ."i  jier  cent  nr  more  of  CO.^.  That  tlii- 
rise  was  due  to  incr<'ase  in  tlic  CO^  tension  was  shown  luit  only  hy  fiiidiirj 
a  irreater  percentasre  of  CO^  (col.  l.'>)  in  the  Idnod.  hut  also  Iiy  heiiiir  alili 
to  demonstrate  that  when  CO_,-free  air  was  liulihled  throusrh  the  liloml 
iciiiii\ed  dnrinir  the  dvspnca,  ('h  immediately  rettirned  to  the  normal 
wliicli  it  also  did  when  tlie  lilood  removed  after  the  animal  had  lireathol 
■fur  a  few  minutes  in  outside  air  (col.  I(ii.  The  CO^  cnnteiit  likewise  n- 
turned  (cnl.  17i.  Had  the  increase  in  acidity  lieen  caused  liy  lionvolatil' 
acids  lactic,  for  exami)le  these  results,  particularly  tiie  latter,  eouh 
not  have  lieeii  nlitained, 

Altliou<;li   there  is  therefore  no  donlit   that    the   ('„  nf  the   Mood    ma 
he  raised  heeause  of  an  increase  in  CO^  in  solution  in  tlie  Mood  ])lasma- 
a  ( '( >^  acidosis,  as  \vc  ma.v  call  it    (see  [laire  .l.'il  I  -   this  does  imt  prove  tli.-i 
the  stimulation  of  the  res]iiratory  <'enter  is  liro\ip:lit   aliout  solely  liy  C, 


Till:    riiNIKul,   (IK    Tin:    Hi  SI'IKXTlnV  .!.>.! 

lilt'  iiicrriisc  ill  till'  i-iii'liiiiiati'  inn  IK'O.  imi  ilscll'  liUKlit  also  scivt'  as 
I  stiliillliis.  That  Mli'li  is  actiiall.N  tlir  casr  was  liciiiciiistratril  jiv 
liiHliiiir  tliat,  if  the  (',,  of  tlic  MikkI  was  tirst  i)f  all  lowcroil  liy  inject iii'_' 
ilkiili  iiitravciioiisly,  livpcipiicii  still  tlrvt'lopcd  in  ]ii'ii])i>i'tiiiii  as  the  t '( > 
.ii'ciiniutati  <1  in  tlic  iiispircil  aii".  ami  tliat  ('i,  nf  tlio  liloinl.  wlien  tlic 
lni>orpiica  XMis  at  its  liijilii-st.  was  hfhnr  tliat  nf  iioriiial  liiood.  Snini' 
nilicr  factor  tliaii  ("h  iiiiist  olivioiisly  lie  icsponsililo  for  tliis  icsiilt.  This 
must  lie  tlie  HCO^  anion. 

Iin    Kkkk.ct  of  Ki:ni!f:.>TiiiN(i  rMuuiN  Dioxnir.  on"  tiik  Mini  tk  Vcpi.imi:  ash  (■%  Tin 

H  ION  roNCKNTKATIiiN   .AMi  Toi  \I.  C'aILHOVATK  CoNTIM    OK   Till    AKTIKIAI. 

Ili.O'ii)  IN   Mil:  |)K(i;iii:ni!ATK  Tat 


I'KILIMINAin    I'CKIOh' 


KIIIIII.ATIfIN*.  ri  lilnli 


HITKKN 
MIMTKs 

A>-reii  HF.- 

HHf  A1MI\<. 
riKlnli 


I  ! 


Time 


a    I  1- 


•■■■S 


<*  —      n 


-,tg 


kH 

-'' 

:i6 

2.2  10.:«la.m. 

:w.5  4« 

■>H 

33 

2.2     l.ri()  |i.m. 

:«.«  22 

.'9 

35 

2.(1   I().:«)  p.m. 

3(<.6  28 

:;(! 

;J2 

2.3   11,00  p.m. 

:w.7  32 

:H 

31 

2.>)  11.15  a.m. 

m.H  32 

8(>1 
HI6 

7»W 
«96 


7. 15 ; ;«).:» 
7.tO  40.2 
7.40  I  :w.i 
7.45 :  30.2 
7.40   32.0 


ertil 

M  3,96H540 

40  2,KWi:t«7 

411  3.>a0  3K9 

54  4.53H4«9 

60  4.MN)43H 


;ii-r 

6.45 
5.02 
5.35 
6.40 
5.95 


7.25 
7.20 


53.  t 
5K.0 
51.1 
51.1 
.50.0 


M 
r.35 

r.4 
r.4 

M 


:tt.2 
12.0 
lii.o 
:t5.ii 
352 


Avt-rage 


:<2.l  7M5    7.42    34.1 


"I'm    II  tllp  aitvi.ll  valiir  Riven  in  tlic  tatilr.     Tl'i 


riv  |«rii].nrtinn,Tl    !•<   t'li 


Further  corroboration  of  the  claim  that  the  UVO.  anion  has  a  si)i>cific 
^tiiiiulatiii!,'  elTect  on  the  respiiatory  eoiitcr  that  is  iiulepeiitlent  of  ("n. 
iiiis  lieen  furnislietl  liy  l[ooker,  Wilson  and  Connett.-'  Those  authors 
^lll'C^edpd  in  keoiiinj:  the  centers  of  the  medulla  alive  liy  perfusion  Mltli 
'letilii'inatod  lilood  tliroufili  tlie  Mood  vessels  of  the  hrain,  and  found 
I  hat.  although  the  respiratory  iiioveiiieiits  of  the  tliaphragm  became  de- 
I'lf.sscd  with  a  decrease  iiiul  excited  with  an  increase  in  ('„  of  the  per- 
I'lised  fluid,  a  Rieater  activity  of  tlie  center  was  produced  when  tliis  coii- 
!ained  a  hifih  tension  of  ('()_.  tliaii  with  another  tluid  of  the  same  (',, 
liut  with  a  low  tension  of  ('()_,.  HV  conclude  that,  althotinh  thr  T,.  i.s-  ihr 
important  rcspiraiorif  hormatic,  the  carhoiiatr  ion  [IICO.  anion)  also  has 
•I  .itimulating  influence. 


i  mr\k 


\i 


«      t 


ii  1 


iH 


'  m 


'■' 

:i| 

1 

■■    ' 

1  ■  i 

'  ■       "1 

I     ■£■ 

■    <                   i 

If 

;! 

Pl 

\l[ 

\ 

:\:,\ 


Till;    IIISIMII\T"iV 


Relationship  Among  Acidosis  Conditions,  Alveolar  CO,  and  Respir 
atory  Activity.     It  wiW  \<i'  \>\i\\\\  timi  vmiiitions  in  tlif  rcspinitnry  lim 
inline,   w  liiitcx  cr   this   may   lie.   iirc  iissoriiitcil   witli   cliiiiiitt's   in   tlic  Co 
I'liiilciit    III'   llic   iil\  t'liliif  iiif.     t'losci'   I'Miiiiiiinl  inn    luis   shown,   Imwt'vci 
that  this  ri'hil  iiiiishi|i  is  liy  im  iiicaiis  always  so  siin|ih'  iim  in  tin-  iiiNtanri^ 
just  (h'scrilii'il,  w  hi'ic  incn-asi'il   ifspiialion   was  t'ninnl   to  lie  assnciatril 
with  an  iniTcaM'  if  aUfnlnf  ('<•..     Tlicrt'  arc  many  cases  wliere  the  re 
verse    feliiti<pnslii|)    (ilitains     namely,    deeicased    alveolar    CO,    ami    h> 
perpnea.     The  wliole  ipiestion  is  vei'y  elosely  linkeil  with  that  of  the  eon 
Irol  ot'  the  reaction  of  tlie  lioiiy  lluiils  anil  with  the  etiolmfie  factors  in 
aeiiliisis.     When  it  is  t'ully  aiiswereil,  many  oliseure  clinical  comlitions  in 
w  hich  lespiratory  ilistnrhanees  ociMir  will  lie  mnch  hcttcr  nmlerstooil  than 
they  are  at    present.     On  accimnl   o|'  its  irreat    impoifance,  consiilci'ahli 
attention  will  lie  ilcMiteij  in  the  nc\t   few    patrcs  to  soiiu"  ot"  tlie  researches 
which   have  liccn   iiiaile  lic;i?in(r  on  the  relationship  lietwecn   the  CO.  nl 
the  jilveolar  ail'  aiul  the  various  niodilicil  types  of  Incathini;  that  can  he 
proiluccil  experimentally    or  liecome  liexclopcd  umler  altereil  pliysiolonie 
comlitions. 

We  shall  considei'  these  conditions  in  the  follow  iiii;  order;  (1)  Con 
staney  of  the  al\col,ii-  CO,  inider  noiinal  conditions  and  during  niinleratc 
\ariations  in  haromeiric  pressure.  (2i  The  ipnintitalivc  relationship 
hctwcen  an  ;irtilieiall,\'  induced  increase  in  ;dveolar  CO.  tension  fas  li.\ 
liiiathiiiir  CO, rich  aii'  and  tlie  incrciised  respiration,  i.'t)  The  results 
of  these  olisci-valions  will  demonstrate  ii  very  jji-ecise  relationship  to  exist 
lietw"en  alveolar  CO^  tension  ami  respiratinn,  Init  if  we  pioceed  to  lepeal 
the  hitter  olisiiv  jitions  under  conditions  wlteie  the  ;iccumulatiiin  of  CO 
in  the  inspired  air  is  acconi|)aiiied  hy  oxyj,'en  ileticicncy  (as  hy  lireathiii'j 
in  a  confined  space  i,  we  shall  see  that  the  rel;itionsiiip  no  loiiLrer  holds, 
indicatin;!  that  tlic  oxyecn  deficiency  has  caused  somethini.'  to  happen 
w  liich  disturlis  it. 

We  shall  linil  that  the  disturliiiii;  factoi'  is  accumulation  of  unoxidized 

acids  in  the  hlooil.  and  this  will  naturally  lead  us  to  study  tl onditinns 

in    which   sudi   .Mcids   miirlit    develop:    namely.    (4l    Hreathini;'  in    rarefied 
air  (iMiiuntain  sickness  1.     ( .'>  i    Apnea.     ( (i i  .Muscular  exercise. 

In  succcciiini;  chapters,  wc  intend  to  review  the  vvoik  in  these  fields  in 
consideralile  dcl.Mil.  jiarlly  liecause  of  its  \eiy  important  licaiin>r  on  the 
'jeiieial  i|ueslion  of  the  control  of  the  respiratory  center  and  |)artlv  he 
cause  of  the  liirht  the  oliserv  ations  throw  on  the  natuie  of  the  mcchanisiii 
involved  in  the  adjustment  of  the  (',,  of  the  lilood  and  (issues. 

As  we  have  seen,  much  work  com-erninLr  the  physicoclu'mieal  ])riuei]ilcs 
involved  in  the  contiol  of  the  reaction  of  the  lilood  has  lieen  contrihutcil 
diiriny:  recent  years  hy  jihysical  ami  liio|ii}>icid  chemists,  hut  much  of  this 


Till     iiiMHiil,   ftK   nil     IIIMIIMIHiV 


:W 


work  ill  tiiir  jiiiluiiu'iit  I'liils  Id  |iiiy  siitliciciit  icminl  to  llic  rxtiiiKi'tliiini  il\ 
(iiiiiplicalcil  ciiiKliliiiiis  cxiMtiiiK  in  tin-  aiiiiiiiil  lioil.v,  tin<i  iiioii-  pHitiiMi 
l.irly,  to  coiivl.itc  the  |>iiit1v    |ili.vsin>clii'iiiiciil   iliilii   uitli   Ilic   iiiiiiifioiiN 


ilisci'viitioiis  tliiit  tiMvc  rniiii  tiiiK'  to  lime 


"'I'll   r I'ljcil  liy   |)l!\  <<loloi;ist> 


•l< 


■u'liitlintf  tlic  liflinx  ior  ol'  tlic  icspiijitoiy  ci'iitfi-.     I'liv  ^ical  (lii'iiiisls  have 


iitly,  for  cxaiiiplf.  uoiit-  so  I'ar  as  to  ilt-lit 


Ml-  ariilosis  as  a  <'oii<lltloii   i 


I  liirli  tlit'i'i'  is  a  iliiiiiiiiitioii  in  tlic  Itirarlioiiati'  roiiti'iil  of  tlic  lilooii 


III 


iliii'i'd  liv  till*  iliscli 


arifc  into  it  of  (ixi-il  ariiU.     This  is  uoinif  too  far,  f< 


it  fails  to  n'coitnizt'  aciilosis  diU"  to  an  incrfasf  in  tlic  ('(»,  of  the  Itlood. 


Il  is  l)i«'  niolccnlar  rati< 


L.VaHCoJ 


•^  I    .vllicli  (It'tfllllilics  till'  tflisioh  of  to, 


Wlicii  ('0_,  is  adilfil  to  file  lilooil,  fillicr  ('xpfiiiiicntally  li.\  icsi)ii  iiiif  tin- 
1.MS,  or  iiattii'iilly  owiiitr  to  iiiii.sfiilar  *'xcr<-is«'  or  to  patliolo)rii'  i-oinlitions  in 
uliicli  there  is  a  ilefieient  exi-refion  of  CO,,  as  in  heart  disease,  the  ten 
ih'iie.N  of  the  e(|iialioii  to  ehaiiue,  hy  increase  of  liie  iiiiliielator.  is  pre 
\eiited  partly  liy  stiiiiiilation  of  «he  respiratory  eeiiter,  uhieii  u'f'ts  rid  of 
<<>^,  and  jjiirtly  hy  an  increase  in  the  deiioininalor.  The  respiratory 
center  is  so  sensitive  to  sli"ht  increases  in  ('i,  I  hat  it  hi'conies  excitcii 
I'cforc  H  suflicieni  increase  in  IL('<>,  has  occurred  to  disturb  tlic  o"  nial 


I, it  10 


LNallCCvJ 


When  fixed  ai-ids  are  added  to  the  hlood  tin 


iiiator  of  the  e<|uation,  NalICO,,  is  lowered  and  consecpieiitly  ('u  rises, 
•iiid  increased  resjiiration  oeciirs,  lowering:  II. CO,  and  thus  reestalilisliini; 
I  lie  ratio. 


I'3 


1-3 


(II.M'TKK  XIJ 


■1         1 


f}.i 


TIIK  CONTHOI,  OF  KKSlMHATIo\  iContMi 

THE  CONSTANCY  OF  THE  ALVEOLAR  00,  TENSION 
UNDER  NORMAL  CONDITIONS 

Siiiof  II  cliiM-  rclii, ioiiship  cxIhIh  lirfwri'ii  llic  iilvi'iiliir  to,  ti'iixiiin  anil 
the  ri'Miiii-iitiin    iirtivitv,   it   is  to  lie  «•    'cctrd  tlint   llic  tun  wixild  licar  -.i 
strict  proportionnlity  to  I'licli  other,  uiid  since  the  l>rcatliii:u'  under  iioriiiai 
coiiditioiis  does  not  vni.v  niueli,  tlie  CO,  tension  slio\iId  also  lie  eonstaiii 
.Many  oltservations  show  this  to  he  tlie  case.    The  tension  is  reninrkal>l,\ 
I'onstant  fioni  day  to  da.\-  and  even  t'loin  inontli  to  month  in  the  sani' 
indixidnal,  jtiox  ided  tlie  physiolosic  conditions  are  the  snme.     A  sli'.'li' 
seasonal  variation  is  said  to  exist,  a  rise  in  the  tcniperntnre  of  the  en 
vironnient  of  the  individnal  cansintr  a  sli!.'ht  depiosion  in  the  CO,  ten 
sion,  uliile  a  fall  in  tcin|icratnre  causes  a  sli^'ht  rise  (llaldane).     TIicm' 
chantfcs  are  ind<-|)cndent  of  any  denionstralile  chanKO  in  rectal  temper 
atiire  and.  therefore,  are  prolialily  dne  to  the  intlnence  of  the  teniperatnri' 
on  the  '•kin 

>!inee  it  is  the  nninlier  of  moleciiles  of  ('(».  in  a  iriven  volume  of  alvc 
olar  air    i  ■• ,  tiie  partial  pressine  or  tension  i   that  is  of  importance,  it 
is  only  when  tin'  luiionietrie  ]ires>nre  is  the  same  that  the  pirciiihuir  of 
<■<>,  in  the  samivie  of  alveolar  ;iir  <aii  l>e  constant.     To  allow  for  this 
all  resnlts  are  reduced  to  stanilard  harometric  (iressnrc  ("fiO  mm.  Hir 
If  tl..    liarometric  pressure  is  lowered,  there  will   have  to  he  a  hiprher 
percentasre  of  CO,  in  the  sample  in  order  that  there  may  he.  the  sanp 
tension  of  this  fras  in  the  air  of  the  alveoli;  and  vui  vrrsn  when  the  liar 
ometric   j)ressnre   is   raised.     The   e(|iiation    by   which   this   tension,   c\' 
pressed  in  millimeters  of  nu'rcniy,  is  determined  is:    100:7()fl::a:p,  where 
a  is  the  peicentaKe  actually  found  in  the  air  of  the  samplinsr  tulie  and  /> 
the  tension.     A  correction  must  also  he  introduced  in  this  cf|Uation  1" 
allow  <'or  the  vapor  tension  <if  the  air  in  the  alveoli,  for  of  course  H..O 
molecules  will  hehave  like  CO,  molecules  in  causing  a  partial  pressure 

When  reduced  to  this  standard,  it  has  hcen  found  that  the  tension  oi 
CO,  in  the  alvco-i.r  air  remains  constant  under  the  difTerent  harometiii- 
conditions  that  olitain  at  the  toj)  of  a  mountain  or  at  the  foot  of  a  dce|' 
niiiie.     This  is  shown  in  the  followinjj  table: 


'tun   ((INTKIH,   UK    llll.    HI>l'll<Ari(>: 


;j57 


(1) 


ll.\Rl)Mr.TKIi 


I  MM.  IKi  1 


ri>,  MriAi.i  V  roi'Mi 

IN  llllV  AI.VKII   Ml 


( i'i.»  f'KN  r ) 


Tup  of  lien  \i\i«  tUtl,."( 

oifiii'i  ::i-i 

K...it  ,.f  |)„1.m:,i|i  Mill..     s::j 

('"IMpri')4«l'il  ;ii|    .•l.lluil       Ijlill 
■rill    l.giirr.  ill  llin  ..jliimn  art  «iri»i.|  ..I  l.v  ihr  ("imiila 


6.H3 
3.M 
8.S1> 
n.3L* 

H-j  A  .  V 


f.T) 
rMirtxt,  I'HKMHi'Rr 

lift  II,  IN  MniMT 
M.VKOI.AK  AIK  AITtK 

•  Ai.f'i'i.ATiso  ror. 

HAKOMK.TBlr  HHKHNrKI 

a.2a* 

S.8.t 

s.m 

l\  wlirtl   I*         fjnurr*  in 


..|   .iliimii;    II'  I'luurr*  in   (Im  ml, hum;    A         •'lur'i'i'.   li-,i»i"ii   nf  aUinlar  «iri   I''  fifiirrt  uf 

.  .n.l  i.iliimn;  B        h.irnni.lric  |.risMii,    .il   ..a  Ifvrl      A  i>  nl.laituil   (rnin  t.il.lit  gitiiig  llir  «'|Mfoin 
«i"n  at  ilmrrpiit  ti  tn]>rraturt»«. 

('Iijinu'cs  ill   tlic    ric(|iiciicy  of  lii'i'iltliiiii;   that    aif   within   physidKitfic 

iiiiits  liaxf  Mil  iiilliifii Ill  tin-  ti-nsion  nf  aUcolar  CO,,   provided  that 

"Aactly  th«'  siiiiic   tiiiic   is  laki-ii   in   pcrformiiii;   thf   t'ori'oil   cxpiratioiis 
•  luriiiif  which  tlii'  saiiiph's  of   ilvcnlai-  air  for  unnlysis  arc  removed. 

The  Degree  of  Sensitivity  of  <he  Respiratory  Center  to  Ohangfes  in  the 
CO,  Tension  of  the  Alveolar  Air 

This  cMii  lie  d*'ti'i'iiiiiicd  liy  oliscrviiiir  tlic  allciiilioiis  piiMliiccd  in  thi' 
.Illume  of  air  that  actually  enters  the  alveoli  falveolar  ventilation)  dnr- 
M'.'  lireathiiic  in  atiiiosphercs  eoiitainiii<:  ilifferent  iiereentaRes  of  ('0.^. 
Ill  man  an  incii'ase  of  from  O.'J  to  ().:'.  jier  cent  in  the  alveolar  TOj  is 
^'itlieiriit  to  doiilile  appro\imalel\  the  alveoljir  ventilation;  or,  more  pre- 
1  isely.  jiii  increase  of  fen  liters  in  the  ,iir  enterinir  and  leavinir  tlie  alve- 
"li  i)er  miniifo  is  caused  by  raisinir  (he  .ihcolar  ('(>.  tension  liy  from  2.2 
!'i  :l  1  mm.  Hi,'  fDoiiirlas,  ete.'i^'. 

THE  ALVEOLAR  CO    TENSION  DURING  BREATHING  IN  A 
CONFINED  SPACE 

We  have  already  eniployed  similar  cNperiments  in  asccrtainiiifj  whether 
<  <>,  aei'iimulation  or  (•,,  (h'pletion  is  rcsitonsihlc  for  the  hypcrpnea  pro- 
iiiecd  under  these  conditions.  \Vc  concluded  for  the  former,  hut  now 
•II  closer  examination  we  shall  see  that,  althousrh  our  conclusion  was 
"irect,  the  deficiency  in  O,  also  has  an  indirect  efTect  on  the  respiratory 
'liter.  This  is  revealed  by  the  fact  that  thr  Icnsioi)  of  thr  CO,  in  tin 
'h-iiihiv  air  does  nol  incrraxr  in  proportion  to  the  ohscrved  incrcaxe  in 
i'>ihnonar]i  ventilation.     We  must  condud'.'  that  the  'lecrense  in  Oj  has 

c  ciTcct.     l[ow    may  this  lie  explained?     Two  possibilities  exist:  (1> 

■iiat  the  O.  want  has  caused  ortfauie  acids  to  aecumulale  in   the  blood 
Hid  so  raise  the  (',i ;  and  (2"!  thst   in  the  absence  of  »  certain  tension  of 


i  1 


■m 


i  I 


;jr.8 


iiii;  Ki:si'in.\TiiiN 


i 


p? 


liL 


1:  I, 

I. 

i 

'  i 

! 

;, 

■ 

i  Mm 

W 

ii . 

«»,  till'  cxcitiiliility  of  the  i-ciitcr  is  niisfd  (i.e..  its  "tliroslioM"  lowered 
so  tliiit  it  lieeonies  stiiiiiiliited  liy  ('n,  to  wliicli  ordinnrily  it  does  not  n 
spoiid.      We   sliidi    now    proeeed    to   i-xiiiiiilie   tlie    ex|)ei'iiiieiitid    evideiu-i 
liejifiiifi  on  tiiese  )iossiliilities, 

r.y  exiiniiiiatinii  i,\'  tlie  iilveoliii-  iiir  of  ;im  iiidi\  idnid  conlined  in  ii  pneii 
niiilie  ealiinet  in  wliieli  tlie  liiironu'tiie  ])ressiiie  is  iri'Mdunlly  lowcrcil. 
it   liiis  iieen    found   thiit   altiiout:li   tlie  CO.   tension    ceiniiins  eonstiint    foi 

'•'  siderahle  r.iiitre   (cf.  paire  :!.')()).  it   iie^riiis  to  fall   when  tlie  I>aroiiiet  ij,- 

|>ressm-e  lias  reaclieil  ahoiit  .").")()  mm.  Hj:.  At  this  pressure  the  teiisi<iii 
of  (>,  in  the  ai\eoiai-  air  will  lie  {>2  mm.  iiisteiid  of  its  noinial  of  alioiit 
Kl.'i  at  atmospherie  pi'essuie.     I'.elow   it  the  alveolar  i'il.  tension  (|uiekl,\ 

falls,   and   a1    the   same   time   liypei|)nea    li< nn's   evident.    altholl!;li   tlii 

(leison  himself  max  lie  uii  iware  that  he  is  lii-eatliin};  more  dee])ly.  It 
this  experinn'iit  is  rejieated  with  the  ditl'ereiiee  that,  as  the  pressure  i^ 
hiwerod,  an  exei'ss  of  O.  is  introduced  into  the  ehamhei-,  tlie  hyperi)nea 
does  not  supervene  until  a  liaronn'trie  [iressure  has  lieen  I'eaelied  that  i-- 
distiiictl.\  lower  than  when  no  excess  of  ( »^  is  ])i'esent,  and  at  the  sanii 
time  the  < '( » ,  tension  in  the  alveolar  air  remains  iniehan^red.  The  ex 
planation  of  this  result  is  that  liy  lowerin;,'  the  ( >^  tension  in  the  alveolar 
air  and.  therefore,  in  the  Mood  and  tissues,  oxidative  processes  hecoim' 
de))ressed  so  that  inioxidize<|  acids,  such  as  lactic,  accumulate  in  tin 
lilood  and  liy  addiufr  their  efl"cct  to  that  of  the  CO..  sei-ve  to  fiiise  the  (',, 

of  the  lilood.     As  a   result,   the   rcspiratorv    center  Ik iiu  s  excited,   hy 

Jierjinea  supeixfiies,  and  tin-  volatile  CO.,  is  removed  from  the  lilood  intn 
the  alveol.ir  air.  On  supplying  O.,  artifii-ially.  this  failure  of  i)ropei 
oxidation  does  not  set  in  and  Ini-athiii!,'  y^oes  on  iiormallv. 

There  should  he  a  sta)ic  in  the  aliove  experiment  durinjr  which  tin 
CO,  tension  of  the  alveolar  air  is  Inm  usi  ,1  naiiu'l.v,  when  the  fixed  acids 
first  appear  and  decom|)(ise  the  carlionates  of  the  lilood.  This  staire  has. 
liowe\er,  not  Iieen  detected.  When  a  person  is  kept  in  such  a  chamlic' 
for  some  time  at  a  iiressure  which  causes  a  diniinution  in  the  alveohn 
Ctt,  tension,  the  tension  does  not  immediate!.\  return  to  its  normal  level 
when  atnnispherie  air  is  ai:ain  lireathed.  indicatiiitr  that  the  fixed  aciiK 
are  only  slowly  irot  rid  of. 

'I"he  s id    hypothesis     namely,   that    llie  O,   deliciciic.x    directly    raisin 

the  cxcilaliil:;  i  oi'  the  respiiator.v  ccnti  r  has  man,\  advocati's.  aiiioic. 
tlieiii  Lindliaiil.-  who  fotnid  that,  when  the  percenta<j:e  of  O,  in  the  ahc 
olai-  air  was  raised,  a  hiirher  pcrc<'nta<ic  of  CO_,  x\as  lU'cessary  to  eaiiM 
an  incrcas  in  tlii'  ventilation  of  the  hnij-s,  and  eon\ersely,  that  a  distinci 
inereaso  in  'he  excitjdiility  of  the  center  occurred  when  tlie  insjiired  aii 
contained  less  than  the  normal  iicrceiitayc  of  ( • ,,  Altliouu'li  it  is  ,'id 
mitteil  liy  I'.aldanc  and  his  school  that  such  alteralioii  in  the  excitaliilit,\ 


'iiii:  cuNTKor.  iiy  Tiii:  uksi'ihatihn 


:ir>!l 


(iT  llic  iTspiiiiloiy  cciitiT  III  the  <  h   dI'  tlii'   IiIihki   iiiiiv  (ii-iMir  Jit'lcr  luii!;- 

riiiitiiiiicil  cNpiisiirc  III'  til liter  to  tlic  cliiiniiod  tcisiiiii  (if  <•_,,  yet  they 

ilriiy  that  Mirli  iiltcratiiiM  can  (icciir  as  a  tciiiiiiirai'y  (•(iiiditinii.  Tlit'si- 
\Mii'kt'rs  I'diiinl  that,  in  (H-dci-  to  raise  tlie  )>uliniiMic  veiitihitiiiii  hy  10(1 
pel'  cent,  tlic  i  lie  I  case  in  the  al\  eohif  <  '<  >;  teiisinii  re(|iiiie(l  w  as  piactieally 
tlie  same  ( 0..'{  per  cent  i    wlieii   the  iiispired  air  eoiitaiiied  "id  per  cent   nt' 


( I,  as  \\  lien    it    eiiiitained   .">} 


per  cent. 


in  tiie  i)hser\  atiiins  already  lel'eiicd  to  on  the  (h'eereliiiite  eat,  \{.  ^V. 
Scott"  has  secMied  some  evidence  tiiat  would  seem  to  support  llaldaue's 
rdiileiitioii.     Me  found  that  the  response  of  the  respiiatory  center  to  the 


liciceiitaue  ot  (  (I  in  tlie  respiieil  air  was  exactl\  the  same  whether  the 
l.itlcr  conlaiiieil  a  low  i  1;!  I4i  or  a  hiu'li  !•'!<)  and  o\ cr  i  pereeiitajre  of  (>.. 
The  possiliility  that  the  e\citaliility  o.  he  respirator^'  center  is  atTeeted 
dii-eetly  liy  the  (t.  tension  is  to  lie  considered  as  one  of  the  most  iin- 
poitaiit  jirolilems  awaitiiii:  solution. 

Kveii  if  it  iiia.\  have  a  ei'rtain  inlluenee  on  the  excitaliility  of  tlie  le- 
-■liiiatory  ccntei-,  ( )..  deticieiicy  /.■(/■  m  can  serxe  as  a  din-ct  stimulus  of 
the  center  only  when  it  is  of  extreme  deirrce.  Much  lijiht  has  lieeii 
iliiowii  oil  the  ri'latioiiship  of  (».  to  respiratory  activity  liy  oliservinH: 
llic  respirations  diiriiiir  lireathiiic:  in  and  out  of  rulilier  ltay;s  tliroufrli 
■-'Ilia  lime  alisorptioii  Imttles  of  sufficient  size  to  remove  tlie  ('('^.  We 
lia\('  already  seen  that  even  the  ireneral  I'esults  of  such  observations 
patre  :!4!h  show  dcai-ly  how  much  ni'Mc  potent  a  respiratory  stimulant 
i>  accumulation  of  CO.  than  deticieiicy  of  (),,  .More  particular  investi- 
'jatioii  in  which  the  aKcolar  air  is  analyzed  hears  out  these  eoneliisioiis 
.iiiil  at  the  same  time  indii-ates  the  exact  conditions  under  whieli  oi's^anie 
.niiis  heeome  dexclopcd. 

With  a  \  cry  small  ha'j'  'a  lew  lileis'  capai'ityi  hyperpneil  of  a  dis- 
ircssiiiLT  type  lint  without  I'vanosis  superxcncs  in  a  few  iiiimites,  and  the 
ilveolar  air  eoiitaiiis  jierhaps  as  low  a>  (i  per  cent  (».  and  4  i>er  eoiit  CO.. 
of  still  ijrealer  interest  and  siiriiiticance.  however,  is  the  t'act  that  the 
I'.iliu  lietween  the  Volume  of  CO,  excreted  and  of  <>,  alisorlied  I  respira- 
'■>i\  ipiotienl  !  dnriiit;  the  hyperpiiea  is  raised  consideralily  aliove  unity, 
Midicatiliiz  that  an  excessive  excretion  of  CO.  must  he  oceurrini,'.  This 
icsult  is  explained  liy  assumiiiL;  that  the  deprivation  of  O.  causes  larjro 
Miiantities  of  lixecl  acids  to  he  produced,  and  that  these  expel  CO^  from 
'lie  lilood  more  (piicUly  tlian  the  O.  is  alisorhed.  In  cnrrolioratioii  of 
'liis  explanation,  it  has  lieen  oliser\ed  that,  after  outsiile  air  is  lireatheil 
i"i-  some  time  follcnviiiu:  the  ahoxc  experiment,  the  respirator\  (luotielit 
iiccoiiies  very  Imr,  so  that  CO.  must  now  lie  accumulatiiifj:  in  the  lilood. 
1 1  the  ;diove  experiment  is  iepc;itcd  with  a  liiri/i  r  hair  laliout  2tM> 
:iicrs.,  so  that   the  o    falls  slowly,  the  lireathiiiLr  can  he  maintained   for 


;  ir 


'   i; 


« 


360 


Till;    Ki:si'lKATl()\ 


a  iniicli  lonjrcr  period  without  (in;)  tvkhiit  si/niptums  of  Intpcrpnca,  ev.i. 
tlioU!.'li  tlif  (),  jMTfPiifiij.'t'  ill  the  alveolar  air  may  fall  as  low  as  in  tli. 
previous  cxperiineiit.  and  tiiore  are  marke.l  symptoms  of  O.j  want,  sue! 
as  eyaiiosis.  twiteliiii};  ol'  the  muscles  of  tlie  hands,  lips,  ete.     The  i' 
spiratory  (|iiotieiif  does  not  lieeome  abnormal  in  this  experiment  indieal 

ii'K  that   1 xpulsion  of  CO,  from  the  hlood  eau  have  oeeuired  as  in 

the  previous  experiment.  Tlii"  eanse  for  the  virtual  ahsenee  of  hyper 
l>nea  in  this  experiment  is  no  dmiht  that  llie  more  gradual  reduction  in 
*y,  of  the  alveolar  air  and  therefore  of  the  l)lood  did  not  hriiid  about  tli. 
aeeumulation  ol'  laetie  aeid  at  a  rate  that  was  srreater  Ihan  that  at  Avliiel 
the  CO,  was  uot  rid  of  into  the  alveolar  air. 


m^ 


!• ; 


'■■]    ! 


BREATHING  IN  RAREFIED  AIR;  MOUNTAIN  SICKNESS 

(n  eonsiderinir  the  ])arl  ])layed  by  fixed  organic  aeid  in  the  eontrol 
ot'  the  C,i  of  the  lilood,  the  most  important  results  have  been  seeureil 
by  oliservatioiis  on  the  condition  of  individuals  Hviii!,'  at  hiirh  altitudes 
As  is  well  known,  under  these  conditions  certain  symptinns  are  likelv 
to  develo)).  the  condition  bein<r  known  as  niountain  sickness.  The  {Treat 
interest  which  physiolo<risfs  have  taken  in  this  sub,ject  has  been  owinu 
not  so  iiiudi  to  the  imi)ortanee  of  the  oi)servations  in  connection  will 
the  c<iniliti(ni  itself,  as  to  the  liu'bt  which  they  throw  on  the  meehanisn. 
of  respiratory  control  and  on  the  cause  for  abnormal  types  of  breathing' 

Moi'c   or   less  hyi)erpiiea.   especially  on   exertion,   soon   ai)pears   in   ii 

rarefied  atmosphere,  and  the  alveolar  CO,  tension  assumes  a  value  i 

sithM'ably  below  the  normal.  For  exami)le,  at  sea  level  the  minute  vol 
lime  of  ;iir  breathed  in  one  individual  was  10,4  liters,  and  the  alveolar 
CO,  lension  :!!».()  mm.  If^'.  After  beiii;,'  some  time  on  I'ike's  IVak,  wher. 
tile  baroiiH'ler  registers  only  4.")!)  miii.  lljr.  l)ouj;la.s-"  found  the  ininiit. 
voliinie  of  air  to  be  14.0  liters,  and  the  alveolar  CO,  tension  27.1  mm.  Ili; 
.\t  first  siirht  the  above  statement  may  seem  to  contradict  one  pre 
vionsly  niaile,  to  the  cfTecf  that  the  alveolar  CO,  tensi(ni  remains  con.staiii 
at  dilTerenl  barometric  ))ressures.  This  ajiplies.  however,  to  the  imnn 
diate  elTects,  whereas  wo  are  now  cfnisiderin;;  the  later  efTeets,  The  im 
portant  jxiint  is:  ITow  arc  we  to  reconcile  with  the  above  liypothesis  tin 
tact  that  a  iliiiiinntiini  in  the  alveolar  CO,  tension  should  be  aeeompanic' 
by  hyi»erpiiea?  .\  solution  of  the  seeiiiiinjr  eoiitradielion  will  not  oiil.> 
bi'  of  inipoitaiH-c  in  connection  with  (uir  ]n'eseiit  iirobleni,  but  will  assi-' 
lis  in  the  iiiM'stiiration  of  the  clinical  conditions  of  liyiieriniea.  in  whicl 
likewise  a  diiiiiiiished  CO,  alveolar  tension  is  often  observed.  Mount()i> 
xirkncsa  ninii  indeal  tt<  roiisiilrnd  nx  nn  intnnirdialr  rnnilitinu  hcttrni 
the  pli  iisiittixiic  anil  tlir  jKithnliifiir. 

From   \^iiat   we  iiave   learned  we  should  expect   the  above  result   to  b' 


!    -II  : 


Ill:  cnsruni.  UK    iiii:  1{i:m'1hatii>\ 


361 


ili'pciuk'iit  ii|)(iii  iiii  inciciiM'  in  tin-  iioiivol.itile  ai'iil  ciinlciit  nf  tlic  lilond 
Tli.it  such  is  i(';ill.\  tlic  cfisc  lijis  l)eeii  eoiiolusively  siiown  lioth  by  titiii- 
lidii  luethods  niul  liy  oIisci'n  intr  tlu;  (lissm-iatioii  curve  oi  hcinogloliiii, 
uliicli.  as  will  lie  cxiihiiiicil  hitcr  i  sec  !)ii<rc  ."iSiii,  iiiiiy  l>c  luaile  to  serve 


a   iji  Xi- 
^^    a»    o 


6EA 
LEVEL 


COLORADO 
SPRINGS. 


COLORADO" 
SPRINGS 


O 
O 

C 

o 

(— 

> 

Co 


00     <o 
o     o 

■'k'-  l-''i.  'riir  huii/i-nf.il  iiittn-iiptc-I  Inns  rtiTt-dil  tin  nuan  i!"i!i;.il  .-dxi^lar  *.<>.•  aii'l  <>. 
Miio  al  s»a  U'vcl  ii.  t  .  iKt-M.l  aii.l  Ncu  Maxt-ii';  the  Ihuk  Iiiif.  alvi'-lar  (."'>.  |,rfsMiif;  ani 
tlllM    lin,-,    aKinlar    O.    |.ii  — iii-.       (l-r..in     ll..ui;la«.    llal.ian,-,     1  l.n.l.  r-.'i,    a.:. I     S.  lui.l.l.i  ,  I 


•  IS  an  index  ol'  tin'  ll-ioii  c(Mieeiilratiiiii  ot'  the  IiIikkI  'I'lic  ex.ii't  cin^iiiieal 
'lalure  nt'  the  ikuix olatile  acids  that  ai-eiiinulatc  in  llu'  lilood  is  imt  as  yil 
^nnuii.     Tun  txpes  of  acid  can  he  llicniirlit  nl',  eiliier  uiinxidi/.eil  urtjanii- 


i     i 


! 


iW;i 


■fif  'Ja 

i 


\    i 


I    i 


i: 


••       i: 


II  i«  H 


.'StiL' 


'I'lii:  m.si-iUAiiiiN 


H 


■f 


I 


rN:i 


■  t 


I!'      i 


jicids,  (if  wliicli   hiclic  ill-ill  iiiMV  lie  tiiki'ii  jis  tlic  ri'iiri'sfiitiitivc,  or  iwu-. 

Uilllic   sMbstillli'i'>.    like   tlic   ill-ill    plliisplliltcs.      Thill    it    is   lint    liii-tii-   ill-ill 
>lii>\vii    hy   l.ntli   (lirii-t    iiiiil    iinrn-i-i-t    t'\  iiit-iii-i'.      Tin-  iliici-t    i-viili-iin-    li, 
l)i'i'ii    t'liniislii-il   liy    UytTrl.    win.    wiis   uniil.lc   to    liiiil    iiiiy    iiicn-iiscd    |mi 
i-(-iitii"_'c  of  tliis  siihstiiiii-c  litlii'i-  in  tlic  iiiiiic  or  in  tin-  lilnoil  of  person- 
ulid  liiiil  lii'i-ii   liviiiir   fill-  sonii'  timi-   in   tin-   fiinioiis   {{ciriiiii    .Miiii,Oii.iit.. 
lint  on  Jlonli-  K'osii.'"     Tin-  indiii-it  iviilcni-c  inis  liccii   funiislicij  l.y  nl> 
si-i-vini.'  till-  timi-  Iluit  it  tiiki-s  iiftcr  tin-  imliv  iilinil  liiis  stiuti-il  In-i-iitliim 
llic  i-iiri-tii'd  iiir  for  tlic  iilvcokir  Cd,  ti-iisim  to  t'iill.  iis  well  jis  liiiii   i, 
i|Uiii-il  to  liriiii.'  iiliout    till-  rci-ovcry  to  tin-   noriiiiil    uln-n   In-  di-si-i-i;ds  i - 
sen  lcv(-l.     Tin-  follow  inn  i-iirvc.  wliicli  is  si-lf-cxpliiiiiitory,  will  illustiiil. 
these  )>(>iiits. 

T'lUs.  on  Pike's  I'eiik.  whi-n-  the  liiironiet  lie  pressure  is  A7>'.)  iiHii.  llu. 
till-  (•<>_.  tension  iit'ter  iin  iiiitiiil  fiill  took  iiliout  seven  diiys  hefore  ii 
(-111111-  to  its  i)erniiiii(-iit  level  for  tluit  Ipiiroiiietrie  in-essure.  iinij  t'oiirteii 
diivs  (-lii|)sed  iifter  ilesi-endiny:  fioni  the  iiioiintain  hefoie  the  seii-leve! 
tension  hiid  lieen  lejiaiiied.  Tin-  slow  iiiituri-  of  these  ehiiimes,  when  i-mii 
piiri-d  with  the  liipid  i-liiiii>:i-s  oliserveil  in  the  i  \pi-rinieiit  with  the  Imii- 
iilri-jiily  iilinded  to  l  piiiie  :;:>S|.  slmirs  ihiirhf  tlial  liiitit  luid  m it  iml  I., 
i-is/toiisibh'  for  tin  iiuTiUKi  in  ll-loii  riiiiri  iitrnlinn  in  nioiinhiiii  ,\(V/,/,, » 
C.y  (-xelusiim  it  would  iippeiir  tliiit  tin-  iin-reiise  in  ('„  is  the  result  of  iin 

(Ni-ess  of  (ixeil  iiior!,'iinie  iii-id   ill  l'(>,i    in  the  hi 1  di-peinh-iit  mi  ii  di- 

proporlioiiiite   exeietion    of   hiiM-^   hy    the    kidneys   duriiiir   the    period    o: 
in-eliiiiiiti/iitioii   to  the   riiretii-d   iiir. 

(Mlier  i.l.servers  iiver  ttiiit    tin-  iii-iilosi>  does  not    leiilly  exist,   hut    tlnil 

the    exeitiiliility    of   the    respiiiitory    eeiiter     itself     In iiies     liiisod     i  it- 

threslmld  lowered',  so  tliiit  it  responds  more  n-iidily  to  the  iioriiiiil  (' 
of  the  hlood.  It  liiis  heeii  ^tilted  thiit  the  iiii-reiise  in  exeitiiliility  of  th. 
i-enter  is  dependent  upon  tin-  jietion  of  the  intense  li;;ht  riiys  iit  hii^l 
altitudes-  the  t-ryllieiri;i  of  the  skin,  ele..  Iieinir  evideiiee  of  this  e\cit 
insr  iii-tioii  of  liuht.  The  eoiis-iint  irritiilioii  of  the  skin,  these  aiithoi- 
siiy.  serves  liy  st iiiiiiliit ion  of  iilVereiit  nerves  to  iiiaintiiiu  ii  hyperexcii 
iiliility  of  the  respiiiitory  eenter,  (Ithels  helieve  lliiit  the  liypi-rexeil 
iihility  of  the  eenter  is  a  direet  result  of  the  iiiiiiiitiiiiied  (>,  delieiene,\ 
The   hiiliinee  of  evideiiee,   however,  stiinds   ill    favor  of  the   view   tliiit    tli- 

P'k'I '••ii''i  "I'  I llitiii,,   sii-kiiis>  depend  on  ehiiiiires  oci-iirrin;:  in  the  in 

or-rilllie    lioli\oliitili>    iiehl-    ..f    the    blood.       The    other    pheliolili-nii    of    till- 
illterestinu'  eolldilion   will   he  discussed   elsewhere    (  piiije   :!|l!l). 


-.1 


'ii     ■ 

•I 


APNEA 


If   a    man    lirciitln-s    forcihly    and    ipiickly    for   about    two    niiiintcs.    In 
will   (-xpcrience  no  desin-  to  bn-iitln-  for  ii    further  period   of  about    tin 


Till;  loNTHoi.  «{■■   riii;  m:si'ii{.\Ti(i\ 


til 


IMC    (liiriitinii      lie    liccdiiM's    ;i|iii(ii".      ^Vll('ll    tin-    ili'siic    to    lnciitlit'    rt'- 
nis,  tlu' lii't'atliiii"'    is  at  t\v>\  vci\  sliallnw,  Iml  iri'inliiallv  ln'cimics  iiinvc 


ii'kfd.  until  at  last  iiorii 


lal  rcspiriil  ;"ii  is  I'lfstaiiiisln'i 


If 


II  saiiipli 


!(>  of 


ilvcdiar  air  is  r('iiiov('<l  at  tlic  tiiiif  wlicii  llic  ilcsiic  tu  IncMtlic  ri'tiiiiis. 


it    will    lit'    i'liiiiiii    to   ciiiitaiii    a    \<'r\    siiimII 
llial  for  SKiiic  tiiiii'  iiicviniis  to  tiir  mi 


|icrc('ii 


ta!,'c   of  ().    iiidicaliii'r 


■t  nf  lifcntliiiii:  there  had  lieeii  in 
llie  alveolar  air,  and   tlierefore  in   the  liloml.  >ip   low    a   pen-entasie  of  (). 


tllM 


t  if  (».  delieieiiev  eoiild  stimulate  lireathiii^,  this  would  ha\e  startei 


•h  earlier  than  it  aetiialK   did.     A 


III  ex|ieriiiieiit  l)>    llaldaiie  is  'jiveii  in   Kiu'.   I-!I.     The  person  may  liejiii 


sliowillii   the   results  of  sUell 


In  show   svinploiiis  o 


f  O ,   want,  snel 


I  as  eyiiiiosis,  liefore  the  desire  to 
lireatlif  returns,  which  furnishes  strontr  proof  lliat  ( >^  wjiiit  itself  ejin 
not  serve  as  a  stimulus  to  the  respiratory  center.  The  failure  of  the 
center  to  act  must  rather  \h;  due  to  the  lowcriiiLr  of  the  ('h  eouserpipiit 
ii])oii  the  removal  of  CO,  from  the  liloml  hy  the  forced  respiration  \vhi(di 
preceded  the  a))ii('a  —  wiisIiIkii  mil  of  lln    CO..  as  it    is  ciilled.     That  this 

has  really   iirred  can    readily  lie   >lio\\n    liy   c-.tiiiiiitintr  the   i'O^   con- 

iciit  of  a  saiiijile  of  alvt'ohir  air  collected  hy  ha\  intr  the  snh.ject  make  a 
'orced  expiration  early  in  apnea.  Kxtrciiicly  low  values  alonij  w  Itli  a 
respiratory  r|uotieiit  (  pa  ire  ."')47)  of  idioii'  (l.'J  arc  often  found,  whereas, 
diiriii;;  the  prcccdiii";  t'orced  hicatliini;  while  the  •'(),  is  lieiiii;  washeil 
out,  the  (piotient    is  often   ten   times  as  ureal      \  iz..  "J.O. 

.\s  would  he  expected,  the  low  O.  perccntaire  iircscnt  in  the  alveolar 
air  toward  the  end  of  the  apneic  pause  is  not  without  some  efl'eet,  indi- 
ici't  thoii^di  it  may  In-,  on  the  excitaiiilitv  of  the  res])iratory  center. 
This  accounts  for  the  tact  thai  the  ;\l\eolar  .lii.  at  the  moment  the  de- 
sire to  lireathc  returns,  usnallx'  contains  a  hiiri  r  |)crcentajre  of  <'(•_.  than 
'  he  normal,  inilicjitiiiu'  that  some  iioiix  ohitile  .-ii-id  iiiiist  have  aceumiilate<l 
ni  the  oriranisni  so  ;is  to  r;iise  the  t'n  of  the  lilood,  and  thus  rctpiirc  a 
lower  tension  ot  i.'().  to  overstep  the  threshold  of  e\cilaliilit\  of  the  re- 
-piratory  center.  In  a<^reeiiient  \vith  this  c\plan;itioii  it  has  lieen  found 
'hat.  if  the  last  tw(i  or  three  forceil  respirations  precedinu'  the  apnea 
.ic  made  in  an  atmosphere  of  (t,  instead  of  air,  so  as  to  till  the  alveoli 

ilh  ().,  the  apnea  c.'in  lie  maintained  t'or  a  very  much  hniirer  period; 
iiid  when  the  initural  desire  to  hreathc  returns,  the  •'(>_.  tension  of  the 
il\colar  air,  iiisiead  of  lieinir  lielow  the  normal,  is  aliove  it.  The  ciTcct 
•f  O^  in  )iidlon!.'inir  apnea  must,  therefore,  he  (lependent  on  the  fact  that 
1  prevents  the  a<'cumiilatioii  in  the  ort;anisni  of  the  luioxidi/ed  acids, 
■  '■avintr  to  CO.  alone  the   function  of  raisiiiL^  the  Cn   in  the  lih-'d  to  the 


.1 


remiired  to  excite  the  respir.itory  ecu 


ter.     r.\    tl 


Us  iiiea'is  the  jierio 


.1 


U   which    the   linath    c 


;in    lie    licli 


I   after    hreal 


lil".:    O,    i 


s   soinctiiiies 


•GU 


Ik 


1 


I    ;i. 


I  ; 


Ill, 


ff'nr' 


;<;i 


Tin;  Ri:siMiiATioN 


■  r 


^ 


i'      t: 


i 


in 

3  » 


liK.  I':  >in\.-  -liiMiiiu  ».iii.in..n-  Hi  .ih,.,|,,i  na5  UiiM.,n>  ..tl,r  t"r>i(]  i,rialhiiiK  f'M  I- 
.mmiU-s.  IImm  1,h,  <),  „„-,,„„;  ihuk  luu-  :-  O)^  n-nMon.  I),„.l.l.-  lim:  .  ,„,rn,a:  alv.n], 
«.<),.  IcnsiM.i,  liolt,.!  luH-  s:iu„s  Ihf  alv.olar  C'O^  tension  at  «liuli  lircathiiiK  w.uil.l  rcc.inni,.! 
..t  til,,  en.i  .,1  ai.ii.,1  uitli  lli,-  alviclai  O..  i.nssiins  sh.m„  l.v  ih,.  il,i..  li,....  riir  a.f.'.al  !T?a!h:- 
I-    ..|.ii.au.l    .,1     lri<     ii.UM-    lail    ,.t     l|.r    Ugmr.       It    is    l..ii.,.|i.     1,,    start     ^.t1ll.       (  T,   ,'„  "  I  ioiiflas"!. 


.   :1 


liljd 


Tlir.   CIINTU^H,   OK    Tin:    HI.SI-linTKtN 


:'.(;:. 


pliciioiiu'iial;  ill  Hill'  iiKliv'uIiial,  for  oxaiii|i1i',  iil'tcr  iiroiitliiii};  lorcihly  I'lH' 
,1  few  niiiiutt's  ami  tlicii  (illiiiK  Ihc  IniisTs  \\itli  ().,,  n|)iii'a  laslccl  I'lir  liijlit 
Miinutcs  and  scvoiili'cii  spconds. 


The  Supposed  Nervous  Element  in  Apnea 

It  is  iioccssai'v  III  point  out  tjiat,  iirior  to  llir  il.ilMiiiilinii  nl'  ni-rinatr 
iiii'tliods  for  tlto  iii\  ('stiifatiiiii  of  tlic  clu'iiiistiy  of  rrs|)iiMtion,  niaii.v 
|iliysiolosrists  iii1fii)rot('d  tlio  ajiiica  t'nllowiiiu;  I'mci'd  ])i'catliiii<i  as  tlif 
irsnlt  of  a  sort  of  inliiiiition  of  tlio  rcspiratiiiy  ci'iiti'r  linniiilit  nhout  )i\ 
its  lopentcd  stiiinilatioM  liy  aflVrcnt  ni'ivmis  inijiiilsi's  transmitted  to  it 
:iliiM<;  the  vajrus  ihtvcs.  tiii'sc  iiiiinilsi's  liiMiiy:  set  up  liy  tlii'  frcfpiont  coi- 
l.ipso  and  distention  of  the  alveoli  ai-tini:  on  tlie  tei'ininatiniis  of  tlie 
nerve.  In  jnstifieation  of  tlie  nervous  iiiter|)retatiiin  of  apnea,  it  was 
ihiiined  Iiy  tlie  earlier  nliservers  that  it  eoiilil  not  readily  he  ])rodiu'e(| 
11  animals  after  se\eiin<i  hoth  \ay:us  nerves,  Mure  reeeiit  work  ha^ 
-Ihiwii  that  tills  is  not  an  accurate  oliservatioii.  for  if  the  seNcriiiLr  of 
tlie  vajri  is  aceoiii]>lishei|  not  hy  eiittiilsr  lint  hy  free/iiiy,  then  a|niea  is 
;iN  readily  produced  as  in  an   intact  animal   i  Milioy  .^^ 

That  ehciiiical  and  not  nervnus  factms  cause  the  apnea  is  fuithei- 
'I'liionstrated  hy  tlie  Mcll-knowii  exiieiiiiient  of  Fiedericii,  who,  after 
li-'atiii;;  the  vertehral  and  one  of  the  carotid  arteries  in  two  dogs,  anas 
t.iiMosed  tlio  central  end  of  the  lemainiii!!:  carotid  of  the  one  to  the 
!"  liplieral  end  of  the  <'arotid  of  the  other  animal,  thus  estahlishini;  a 
missed  circnlation.  lie  th-'ii  found  that  hy  ap])l.\  iiiir  foi'ced  artificial 
r'spiiiition  to  the  one  animal,  the  apnea  \\hich  siiiiervelied  affected  the 
■ 'lier  aniiiial  and  in'!  that  to  whicli  the  artificial  rcspiratiun  IkiiI 
ilMally  hecii  applied.  .\iii«thcr  proof  iif  the  eheiiiical  theory  nl' 
:.;iiiea  is  furnished  h.v  the  oliservatioii  that  if  forced  lireatliinsi  is  iicr 
I'lnied  in  an  atmosiihere  coiitaiiiiiiir  ("(^.  in  ahoiit  the  same  jiartial  i>ies- 
-'iie  as  ill  the  alveolar  air,  no  apnea  supervenes,  ami  if  the  exiieriment 
!■-  repeated  sexcral  times  with  jirofiressively  decliniiiLr  liercentafro  of 
'  ''.  ill  the  air  each  time,  the  leiiyrth  of  the  a]nieic  jiaiisc  proportionally 
i   creases  as  the  CO^  jiressure  in  the  insiiired  air  diminishes. 

.Mthouirh  in  the  l'ore<roinir  account  we  have  ado|ited  Ilaldane's  view 
'-at  oxysreii  deficiency  /ice  xr  can  act  as  an  excitant  of  the  rcsiiirator.\ 
'  liter  only  when  it  is  of  extreme  doKroe.  it  should  nevertheless  he  pointed 
"it  that  studies  hy  A.  S.  LiM'veiiliart  on  the  action  of  i-yaniiles  mi  t! 
I  spiratory  renter  have  led  him  to  conclude  that  interference  with  oxida- 
■'.:<'  processes  may  he  a  more  ]iiitcnt  f.ictor  in  its  stimulation  than  the 

lieiiiiieiits  ill  which  oxyjren-poor  atinosidieres  are  resjiired  wiiuld  lead 
-  to  exiiect.     We  must  await   further  evidence  hefore  a  final  \erdict  i-. 
I'riinoinicod  on  this  most  pei|)lexing  in-ohleni  of  modern  iihysiolofty. 


I     I  ■, 


If 


V.'  j 

i 


i 
fll 


<  i;.\i'Ti;i,'  xi.ii 

tin;  (  u\t|;<i|,  it\'  I.'KSI'IUATION  iC.iitMi 

THE  EFFECT  OF  MUSCULAR  EXERCISE  ON  THE 
RESPIRATION 

l>iliiriu    iiiiiM'iiliir    I'NticiM'    llif    iiiiliiniiiii'    \ ciitilatioii    iiiciciisos    to    :ii 
"•xtijionliiijiiv  extent.     At   rest  iiii  iiveiii!.'"'  iniili   lespiies  tl  to  S  liters  i.i 
iiir  pel-  iiiiimte,  liiit  (liiriii-r  wnlkiiifr  on  the  levi!  ,it  the  nite  of  :.  kilometei. 
.Ill   hour,  this  tiyiiir  IIKI>    ilii'iiMse  to  .iliollt  "JO  liteis. 

The    tilst    investijjiilioiis   iis   to   tlie   ejillse    of   the    leliitiniishij)    hetweeii 
Miiiseiiljii-   iieli\it\    .'iikI    pnliMoiiie    \  eiit  ihil  ion    were    niaile   hy    iiniiiiiil    e\ 
I'eriiiieiits   in   uhicli   tet;iiiiis  of  the   niiiseh^s  of  Hie  liinil   liiiilis  \v;is  Jm-.. 
(Iiieeil  h\    ejeetrie  st  iiiiuhit  ion  oi   the  spiiiiil   eord.     The  proMem  was  t- 
linil  out  uliiit  s(  rves  as  liie  means  of  eorrehitlon  uiei-ve  letlex  or  lioniion. 
eontroli    hetween    the    iiiuseiihn    aetivit.v   ami    the    respiratory   activity 
I'.y  eiittinir  tlie  spinal  cord  ahove  tlu-  point  of  stiinulation.  it   was  foiin.i 
that   tlie  tetanus   was  still   aeeoriipanieil   liy   as  marked  a   hypeipiiea   a^ 
hefore.     On  the  oilier  hand,  when  the  spinal  eord  was  left  intaet  hut  tie 
Mood  vessels  of  the  linili  were  li-rated,   no  hyperpnea   followed  the  teta 
inis.      Kvi(leiitl\    tliei-efore  the  patliwa\    of  eoiiiiiiunieation   is  the  blood 

Tlie  next  step  was  to  seek  in  the  Mood  for  the  suhstanee  or  lioniioiie  thai 
a<'ted  as  the  respiratory  excitant,  and  natuially  the  lirst  |)ossil)ility  eon 
sidered    was   a    ehjinjre    in    the    trases    of   the    lilooij.    either    a    delieieiiev 
of  (>,  or  an   ilierease   in   CO..      Direct   examination   of  tlie  Mood   for  tin 
(piantily  of  these  -jases.  however,  yielded  results  wliieli  were  quite  eon 
trary  to  such  an   hypothesis.     It   was  found  that  the  pereeiifasre  of  O  . 
if  aiiythiiifr,  was  sliuhtly   increased,  and   that    of  the  CO...   if  anythinL' 
diminished.      >Ioreo\er,   when   the   expired   air   was  analyzed   dlirinj;  tip 
hyperpnea,   the  /n  ra  iitdi/i    of  Co,   eiuitained   in   it    was  distinctly   beli'w 
the  normal  average,  ami  the  ))ercenta<je  of  O,  ahove  it.     Kvidently,  thr-  ■ 
fore,  the  niiiiniiit  of  jiases  in  the  Mood  lias  iiothin;:  lo  do  with  the  excita 
lii.n  of  the  r.'spiratory  center,  and  the  conclusion  drawn  hy  the  earlic 
investigators  was  to  the  ctVcct   that   the  e\eitin-r  sulislaneo  carried  frmi. 
the  ai-tive  muscles  to  the  res])iratory  center  must  he  some  unusual  mcta 
liolic  luoduct,  possihly   the   lactic  ;icid   product  d   hy  coiitraetiou. 

It  was  further  found,  by  examination  of  the  resjiiratory  (piotienf,  tlia' 

.■!r,ri 


m 


TIM     (uNIUiil.    Ill'     IMi;    Ul>-ril(\lln\ 


:]f.7 


III  cNiTHs  of  CO.  wiis  lifiiiy  ('X|iirc(l  (liiriiis;  tin-  \\i>ik  ;iiii|  iniiiictliatt'l y 
ilti'i-  it,  liiit  lli;it  this  was  siilisciiiifiitly  t'tilliiufil  liv  t\  iiiucli  lowci'  qiM'- 
iiiiit,  incliciitiiiir  tluit  ('(>,  wns  inMiiir  rt'tiiiiicd.  Such  a  rt'siilt  would  l>r 
ill  r<iiifiiriiiitv  with  ;  \  icw  that  an  arid  siidi  as  hu-tic  is  discharui'd 
ihlo  tilt'  liltMid,  on  the  railiciiiiiti's  nf  uliich  it  wnidd  act  as  explained  on 
|.:iV'<'  :!.'>."i.  I'.rcatliiiijr  in  and  out  of  a  Miiall  iiililicr  Iwiir  causes  tiic  sjiiiic 
Iterations  in  the  i-c>|iiratcii>  ijiiotli'iii   i  see  pajre  .l.'iW }. 

That  hictie  acid  is  aetiiallv  produced  li\  cinitraetini.''  iiiuscje  couhl  not. 
Iiii\\e\er,  lie  shown  hy  all  invest ijralois,  and  it  was  not  until  some  yeai's 
liter  tliat  l'"lctclier  and  llopkins''  clearly  ileiiiinistrated  the  conditions 
'.iider  which   it    may  appear  in   active  isolated   luusele.     These   ohservers 

'iind  that  lactic  acid  is  produced  in  excised  niliscles  onl\  when  the 
iiiiiseiilar  contraction  occurs  in  a  delicieiicy  of  (»,.  When  !t  occurs  in  an 
HJi'ipiate  supply  ot'  (»,  ('(»,  instead  ol'  lactic  acid    is   produced. 

Takiliy  these  laits  |n._'elhei'  with  what  we  alicad.v  know  coiiceriiiui; 
'111'  conditiiiiis  under  wliieh   the   respiratory  center   reacts   to  conditions 

liieli  l)resiiuialil\'  cause  a  change  in  the  C,,  ol'  the  lilood.  we  may  t'oiliiu- 
l.iie  the  hypothesis  tliat  respiratory  activity  diiriiiL'  niuseular  exen-ise 
I-  due  to  a  slijilil  iiiei-ease  in  the  ( '„  of  the  lilood,  and  that  this  increase 
Is  owiiij;  fxirtlfi  ill  nil  iicliiiil  iiiiTiiin(  in  ('(f.  priiihuliiiii  In/  tin  aitlmi 
,'  'isi-hs  (111(1  iKirtlfi  t(i  IIk   iirodiictidn  <//'  Idctlc  arid.     Such  an  hypothesis 

•  mid  satisfiictorily  explain  why  the  ai-tual  aiiiount  of  CO,,  in  the  lilood 
i'ii','ht  he  lielow  the  noriiial  duiiii!.'  muscular  exercise,  for  the  CO,  Axiiuld 
'■  "washed  on'"  from  the  lilood  li>  the  liy])erpnca  induced  l:y  the  in- 
•  "Mse  in  Cii. 

The  olivioiis  method  of  puttinj;  this  hy|>otliesis  to  the  test  is  to  ex- 
Miiine  the  alveolar  ( '< ).  tension  and  the  respiratory  qiiotieut  under  various 
i"itdiii(ins  of  muscular  actiKity.  The  icsiilts  of  such  ohservations  are 
-'i\cii  in  the  accoiiipiiii villi;  talile. 


% 


(  I  1  (-2) 

<),  usi',1  CO,  |ir,i 

ill  I'.r.  iliicfd  ill  I', 

pi'r  iiiiii.  jicr  mill. 

Ullli!!;;  n-l,  >tlU'ilill(;  :'Js  I'lil 
\V:ilUill(;   ;it    til.'     nti-   cit' 

.'I  kijoincti'is  iT  InMii  Tsii  i;i;i; 
U:ilkih;;  al    th      i.it..   Ill' 

."i  kiliiiiictc.^  per  liiiin  I  mi.")  liiJJ 
W.ilkiiij;  ill   tlh'  lilt,'  lit' 

li  kill tcrs  per  liciiii    l.")'.i,')  l::!iS 

\\:ilkiini  lit   till'  lilt"  (if 

7  !.ilii;iuii: ;  per  h.uir  I'fc")  ITss 
W.-ilkilli;  ill    tlic   lilfc  cif 

s  kil'inif'tiMs  per  liiiiii    1'.")  I.'.  :2.",S(i 


( ■•  1 
H.  q. 

Mll.«'(\ 

vTriTT)" 

II. set 
(i.Spl 

ii.siii; 

0.s7t! 
n.s;i| 

n.!i:w 


<''Viii       T.itiilnlvc.lm 

"hi'iilai'      vriitilatiiiii  in 

iiir  liliis  per  mill. 


r,.iii 

I!.  11 1 

(■i.:'.i! 
(;,i:ii 
fi.in 


.I.SII 

I. ■'..(■. 

t^.s 
4 '5.2 


i       rl 


i:ii  ilHi 


:!(iS 


■IMI     111  >l'lll\l'lnV 


III  I  III'  III  >t  I'l  ill  I  III  1 1  i^  uiv  c'li  III!'  ( I.  iiM'il  ill  I'  I-.  I II' I  iiiiiiiiti'.  A  nil  II 1 1;  nil 
tliiii<.'s  tlicsi'  liuuirs  iiiilii-atc  tilt'  iU'lnal  iiiii<>uiit  nl'  wm-U  (Imic.  In  il 
sccniul  riilliniM  is  ^M^l'll  tlir  ('(•.  liriiillH'tiiili  ill  rr.  )»v  iniliult'.  Ily  (li\  i 
iinr  the  lii;iit('s  nf  tlif  sccniiil  cnliiiiin  liy  tliusc  oj'  tlic  til'st,  we  (ilittiili  tl 
liu'iiics  111'  till'  lliiiil  I'liliiiiin.  rrpicst'iiliiin  ilir  I'cspinitiiry  i|Uiitifiit.  'I'l  • 
rtiiirtli  ciiliuiiii  unM-.  ihi-  ('(»,  rmiti'iil  nl'  tlif  iil\  cnliir  air,  aiiil  Ilir  l,i 
I'lillliilli  till-  liilal  (ilcinhir  Ni'iililati'iii  in  lili'fs  pi'i'  iniiiiitc. 

'rakiiii;  fur  llu'  pii'Miil  llii'  liiiUics  in  iln-  liisl  ami  rmiitli  ciilmims  ai 
p(ist|ioiiiiii:  a  I'liii'-iili'iatinii  nf  the  ri"»piratiirv  i|ii(itii'iif,  it  will  he  iml.' 
thai,  as  tlir  iiiusi'ular  wnrk  increases  up  tn  a  total  eiiiisniii])tiiiii  <if  alim, 
1(100  e  1'.  nf  (I  pel-  iiiilillte,  t  lie  i'(K  pereeiitau'e  ill  tlie  alveolar  .1 
sleaijily  iin-reases,  'I'lie  ipieslion  arises,  does  tlie  alveolar  xfiitilali •• 
iiierease  in  proportion  to  tlie  iiierease  in  ('<>,  tension'  If  it  does  -•■ 
inenase  in  ( '( 1  leiisinn  in  tlic  lilooil  ean  lie  lielil  solely  respoiisililc  f^ 
tlie  liyperpiiea      i     e  .  a   piire  ('(I.  aeiilnsis  i ;  Avlienas  if  tlie  li\  |)el'piii'a    1 

trreater  tliaii  ean  lie  ai nnted   for  li.\   {lie  iiierease  in  CO,  tension,  otli' 

ai'ids  must  lie  |>arl!,\    res|)nnsilile  for  llie  aeiilnsis.     I>y  making,'  tins  san 
indixidnal   lireatlie  atiiiosplieres  eniitaininu  ditl'erelit    pereeiitapes  nf  (11 
it    was    fnlliid    tliat    In   prnilllee   a    ilnlllilillir   of   tlie   alvenlar   vetitilatinii 
ni|iliriil  an  inereasi-  aiiiniinliii!.'  to  O  :!:!  per  cent  nf  an  aflimspliere  of  I  '1  ■ 
in    llie   alveolar   air      see   alsn    pau:i'    ■'<■>' K      Wlieii    we    e\aMiini'    tlie   alio 
(iyiiri's  (luriiii:  inilseiilar  e\eieise,  lin\\e\er,  we  find  that  a  rise  in  alvenl.i 

< '( >.  frniii  .">  70  to  (1  :iil  I  i.  e  .  0  till  pi'i lit  i  iniiltiplied  the  normal  alveol 

\ ciitilatioii  liy  eoiisideralily  iiioie  than  fmir  times,  whereas  had  it  In  ■ 
entirely  due  to  an  inerease  in  ('(»..  il  slinnld  lint  have  heen  more  th;- 
twiee  as  iiineli.  I'lx  iiieiitly  therefore,  some  other  faetor  than  CO^  tensi.. 
imist  have  lieen  iispniisilile  I'nr  the  iiieieased  res]>iratoi'y  activity.  'I'l: 
eoiii'lusioii  is  fiiitlier  eonliriiied  liy  exaniitiatinn  of  the  alvenlar  ' '' ' 
diiriiii;  very  streiiiiniis  imisi'iil.ir  ell'ort,  when  a  relative  ilfcmisr  in  t! 
<'<)_,  ]iori'iiita;_'e  lieeoiiies  apparent. 

If  it  is  true  thai  the  exeitiiiu'  ai,'eney  has  lieeii  dei)endent  jiaifly  on  , 
increase  in  the  ('<>_,  tension  of  the  Mood,  and  jiartly  on  the  production 
iioinnlalile   iiri:anic   acids    1  lactic   ai'idt,    we   should    expect    that    imn 
dialely  after  discoiiliiiniii};  the  muscular  exercise  the  ('()^  tension  nf  I' 
alveolar  air  would  fall  to  a  level  distinctly  helow    nnriiial,  that  it  vmmi 
only  slowly  rccii\er  thereafter,  and  that    fiiitlier  exercise  hrfnro  the 
eovery  had  occurred   \Miiild   |)rnduee  only  a   slitrlit    increase   in   alvcn! 
CO..     These  results  we  should  ex|)ec1  liecause  of  the  imich  slower  rate 
which  the  nonvolatile  oriranic  acid  is  i^ot  rid  of  from  the  orfraiiism,  (•• 
jiared  with   the   \olatile  CO^.      i!y  iictilal   experiment    these  supiiositi' 
lia\e  lieeii   fniind  to  he  enrrcct.  as  is  s'lown  in  the  follnwiiij'  tahle. 


}    ,  ?i 


I  III    1  lis  11,'cii    nr    Till    III  -III;  \  iiips 


■  Wi 


I  IMI     \ITI  y.  nisi  IIVI  IM   IM 
\  liniK  I'KKIilli  iiK 

Ml  SI  II  w:  I  \i  III  Ki 


M    I'lii.i.l: 


•.•ii.l   I"iii..,l: 


".i.l    I'liii..!: 


\..l!ii:il    li-li 


1<> 

\-2' 

ill" 
.".li" 

in 

11" 

1" 

.".li" 

.".' 

It" 

s' 

.'!('" 

Is 

:;n" 

■Jl 

i\" 

M  \  1 

M    Ml  1  11     i  1  S-.I 

IS   ^IM.   III. 

.•:"i.  1 

''."i.'* 

:i.*(.>< 

:!«.'» 

.".I.f 

.".llIP 

.it.  I 
:!2.l 

.'■1,1 

"li.'.' 

ll!!  II 

(|)mU:;||.v 


III      tnis      tlllllr      till'      (i^llllrs      III'      I'l'lii.il       1       rr|ilrv|il       lllc      II !  \  I'l  illl  I'      III 

..iiili    in    inlii,    llu    iiiiliirili;ili'l\     I'lilluw  iiiu   ii    |iii  iml    nt'   si  ii'inimis    wmK 

■  lii-lliis    in     Pi'i'iipil    ■_'    nil'    I'm'    llu-    siiiiu'    iiii!i\  nliliil    iil;:iiii     |ii'1  ruiiiiili'J 
siiiiic  iiiiiiiiint   111'  \\iiii<   witli.   Iiin\i'\rr,  milv    ii   slnnl    |iriiiiii   uT  rrst    in 
iiiiii;.''.  .iinl  till-  liiiuirs  111'  llii    tliiiil    |ii'ni  il  iin-  :i    n'|n'1  il  imi  hI'   tin-  s.inii- 

■  liliiiiis.     It   will  lir  iilisi'iv  cij  linit  illl'  iiiiisiMiliir  cxciimsc  nt   liist   raisnl 
.ilvciiljif  ti'iisimi  111'  ('1».  riiijii  till'  iiiii'iinil  iif  :!!•  iimi.  ti>  411. "J  iiiiii  ,  Imt 

'  ill  tlircf  iiiiiiiiti's  iit'tcr  llic  Huik  iunl  iicrii  ilisriiiitinili'il  tin'  ti'iisimi 
~  riiiisi(lfr!il(l,\  lii'liiw  till'  niiriiiiil  I)iii'iii'.'  tin'  si'i-ninl  jii'iinil  uf  mils 
.11-  I'Xt'i'fisc  tile  < '( •    ill  tlic  iii\  riil.'ii-  iiir  I'lijli'i'ti'il  ill 'iliiili'ly  iil'ti'i'  tin- 

■  It   iliil   init    iiK'l'i'jisi'   iiliiiM'   till'   iiiiinnii    Ii'M'I.    iiihI    in   tlir   tliiiil    lii'iinil 
iiii'fiMsi'  w.is  still  li'ss      ri'siilts  wliii'li  an-  I'lilirrlx    in  I'liiiluriiiitx    witli  tlir 

n    lliiit  lis  a  ciiiisi'iiui'iii'i'  III'  tile  (irst    jii'iinil  ill'  iiiiisi'iihii'  I'M'M'isi'  iimi 

iiilr  iiiyaiiii'  ariils  Iiail  ai'i-niiiulatcil  in   tin'  lilnnil,  s,i  ilmt   in   ptml ■ 

1  ri|ilir('il     ri's|iiiatiir>     ai'ti\il>     in     llii'    si'ciinii    ami     lliiiij     iirrimls    a 
.■Il  li'ss  iniTi'asc  ill  t'<>,  iciisiiiii   Was  ii'i|niii'il. 

■\Vi'     lllav      Sinn      lip     ''     '     I'lini'lllsinlls     ulilrll      IItsi'     ulisi-l  \  at  inlls     jllstil'x      iiy 

'iiiL'  tliat  iliiriiiu:  nil,  ■'•iilar  I'Si'irisi-  thr  (',,  uf  tin'  lilnnij  1 iiiii's  sliiililly 

ii'iiscd  licrallsc  111'  till'  lilicratiiili   into  it   nC  ('(•    aini  nt'  lai-tir  ariil   fiinii 

■  ai'tiiiu'  iiiilsi'Ics.      Till'   ri's|'iratiii\    rciilri-  is,   linw  i'\  rr.  sn   ^I'lisitivi'  to 
sliiilitcst    iin-ri'asr   in  r,,   tlial    it    iiniiiriliati'lv    ri's|)iiMi|s  aiiil    iirmiui-rs 

.  liri-jiiii'a,  with  till'  result  that   tin'  \iilalilr  t '< »_  is  su  uaslicij  out   ul'  tin- 

■  ill  that    till'  t'li   is  li.'lil   ijiiw  II   in   spiti'  m'  tin-  i-iiiitinin-il    |iiiiiliii'tiiiii  nf 
'I   siilistaiii'cs   liy    till'    iiilisi-lcs.      'i'hi'    iiinii-   si  ii'iiiniiis   tin'   rxiTi-isc,    thr 

-s  alilc  is  the  <»,  ciiiitcnt  ul'  the  hlouil  tii  ki'cp  jiai-i-  with  tlio  iiK'tahulir 
'ivitx  ul'  ihr  iiii|si-i,'s,  >u  that  li'lat'M'ly  iinirr  anil  iinni'  iai-iii-  ai-lii  i- 
..iliii'i'il,  iii'i-i  ssitaliiii;  tliiTi'tiiir  a  t^ri'ati'f  ami  iiii'atrf  washiii'^  mil 
I'O,. 


1    f 


II 


il  i 


''k,  .MM^ 


.1 


:!7it 


I  Ml     III  •<lll|\TlllV 


ihi 


#'       ' 


hv 


(I 


Th.  i..ji.lihr>,  uiiii  «i,i,.i,  en   ,,.,„  I,..  ,_,.,,  ,.j,,  „,•  ,„,.,,,„,^  ,|„.  |„„„ 

whirU   .A.'it.'N   III.-    i>s|,i,„t„iy   il.-ti\ity    I'lMiu   , timiihi.'   to  lirt   llftiT   it    |. 

II..  Iniii-.f   iv.|iiir..|       l'i..\isi„n    r.„-  III,.   ,,.|i„,\;,|  „f  ;,   Ih.iiii iil'trr  ii 

"■;'''''>    '"'^  ' "  ■li-|'l.i>.'.l   is  ,,r  ,.uins,.  .•ssmtinl   In  ..ni.-init  •irliiti.,, 

"''  •■"""■'i"ll.  iHi.l  is  srcii  ill  III.'  ..,!>..  „r  „|1m.|-  l.nii ,.  s„..|,  „s  .'|.ilirp|| rilN 

.•IMil    sr,.|-,li|i,     uh.is,.    .Iis....lilih|i;ili.'..    ,,f    ii.-li..ll     is    ..iVf.-t.MJ     l,\     ll,,.],.    ,|, 

sinii-linll     III     Ih.'     MimmI      (s,,.     |,i|i,..     74.-,) 

I>ii-...'t   .■xi.l.Mi.M'  tli„t   |;i..|i.-  ji..!.!   Is  l-.„,„,.,|  .lurjiii;  si,    •u.-iis  inns.Mii;,, 

••\.'i.-is..  ill  tniiii   iiiis  I ,1   iiniii>li..|   Ia    Kyir.l    •     IS! ,„„v,..l  f,.„„|  „ 

IHIS..I,   iiMi„..,|i,-,t..v    jiri.T  nniiiiinf  ,it    lull  s| I    i„,-  jilMMit   tliiv.'  iniiiMl.' 

'■'"I'.iiii.MJ  7(i.s  iMiMi-riiiiis  .,!■  Ii.,.ti,.  ,„.i,|  ,„.,.  1 ,.    „,•  1,1, „„|    ,,„.  ,„„.,„.,- 

"' ""  '"•''"-'  '-•''  Niilii..'iiiiiis.     .\|„..|,  ..,•  ,|„,  |,„.,i..  a,.i,|  , umulatin«  i,, 

''"■  '•' I  '"•  "•'  'l"'i'''  !■"'>   li.l  ..|-  l.y  ..xi.lMii.,i,,  l.iii  ;,  |,n,{,>  part  ..r  it   i~ 

ills..  .•X.T..I...I   liy   til.'   iirill.'.    ill   u|ii,.|,    it    «Ms    r.MiM.i    liy    |{ytT..|    h,   ....nsi,!..: 

iilil.'  iiiiiniiiit  iil't.'i'  siifiniiiiis  iiiiis.-iil.'ir  i'\.'iiiiiii. 

'■''"•'">■   ''•'    "^  •>''''•••   I'"'-  ••!    lii.,l.i.'iil    lln    h,l,„rinr  »/    llu    n  sfilnilnr^, 

<l""h,„i.     This  r,iii„  ,is.s  ,,,rl.\    ill  th.    iMiivl,-  vv„ik   (Till.!.-  nii  |.,-,l'.'  :!(i7 
iM.li.-i.finv'  i:.it  1II..IV  CO,  is  l„.iii!r  ...,.,vf,l  ihiH.  (>,  nl.s„rl,.,|.     Aft.T  th.' 
w.,i:.  is  .Iis.-.,ntiiiii.,l,  it  iisnally  fMN  Im.|.,w  ll;,.  iM.rniiil  Imvuiis..  ,.f  ivft-ntinii 

"''  '■"-  '"  '•''^"  ""■  I'l'"' ■  '•"•  ■     '!'•  ii-i.l  thai   is  iM.inir  irra.liiiilly  iis...l  ii,' 

III-   rx.'i.t.'.l,      A    similar    fall    may    s..in  .times   iir    in    ili,.    n.spiratnn 

<|U..fi..|it   il„rl,H,  miis,.|||ai'  ..\..r.-is..,   if  this   is  .•..iitiim...!    t„r  a   h.iijr  tiiii.. 

It  pi-nl.al.ly  i|i.li,.al..s  thiil  a  hah has  hr.,!  stni..|.-  I>..'  ■....„  t!i(>    "f..,lii.. 

ti.m  ..f  la..li..  a.'i.l  in  th..  imis.-l.s  an. I  th.'  h.ss  „(  this  siii.staii.v  l.y  nxi.la 
tion.  Ill  any  .-as.,  it  is  a  si..'iiiti..aiil  ...•.•iiir..|i.-.,  I',,|.  it  .•..iiici.I..s  with  th.' 
Sf.-at   impr..\..|ii..|it   in  th..  siil.j,.,.tiv..  s..|isati..iis  a..|...mpaiiyiii(.'  miis.-tilai 

♦'^•''*''''*''-     ■'  " "'•^-  •'"•■  '■Xiinipl.'.  at  th..  sain..  tiiii<>  as  ih.-  appciran.-i.  ..i 

th<.  "socnn,!  uin.l."  «li..n  ll iivulat..iy  an. I  ivspiiat-.r-y  .listless  cxp.. 

ri<.n....,l  .jiiiini.   th..  ..aili..r  statti.s  „(  sti-..|ii,.,us  imis..iilar"(.x.'i-ti.,ii  .lisap 

r«'.ir.     Till'  sta!i..s  pii..,.  t„  th..  s. M.I  u  in.j  .•.,ri-,.sp..n.|  t,,  th,.  p,.ii.,.I  wh.i. 

.-nsi.l-ial.l..  .|ilalitili.s  .,f   i',-....  co,  a.v   l,..ini.r  u'«i    n<\   «(   iV-.m    th.-   I.I I 

ami  ar..  i.n.lialily  cifatinir  a  tcmiMiiary  iiiata.l.jiislm,.|it  ,,f  the  ('„  whi.'li 

a.'ts  .Ml  III.,  vaii.ius  i jiillaiy  .■..|it..|s.     If  l,y  r.,,..-..,!  I.r,.atliin..'  niu-h  oi 

this  ('<».  is  .Iis,.|iari:..<l  l.,.f.,r..   th..   miis.-iilai-  ..xoivis..  is  Mii.i..ilal<,.|i,  tl,. 
initial   hypcipn.'a   is  imt   n.-arh    s.,  mark...! 


■S 


1       [ 


(  IIAI'TKI,'    M.lll 


Tin:  <  u\Ti{(ii,  di'  i;i:>iMi;.\Tin\   t  ,,i,im 

PERIODIC  BREATHING 

Types  of  Periodic  Breathing 

In  tlic  liot  KiMH\ii  u|'  i!i..>c.  cill.-l  rlii'Mi.' Sl(iki>  if>|.iiiilioii,  i,  piMiml 
"i  liypfrpiicii  >.ii|ii'iM'ii('N  ii|ii>ii  cini'  hI'  iiiiiica,  fjirli  iiciiiHJ  rdllnwinir  in 
iiunJHr  scinu'iii-i',     Altif  iin  ;i|iiiiir  iictinil.  tin'  liicMlliiii!.'  In'.'iiis  jit   lii^i 

liiiritly,  ifiiiiliiiill.v    Ik irir>  iiiiHc  |ii'iiiiiiuiii'i'i|  until   it    is   iiMikiilly  i'\iiu 

'.'I'liiffl,  mill  then  I'inlfs  otT  iij;iiin  tu  llic  iipiirii-  iiiiiiM'.  SnniciiiiM's  tlir 
ii|>ni'ir  pfriiid  is  iinini'iliiiti'lv  t'ollnwrcl  l>\  one  n\'  inti'iisi'  h\  pet  piicii,  tluMf 
lirinir  III)  irffuliiiil  iniMciisi-  In  tin-  icspi,  jitmv  iniiM'ini'iits.  liftwri'ii  tliosi- 
tun  typi's  Jill  \iiliclii's  lit'  till-  ciinilitinli  iili'  liirt   '  l"it;.   I'l'J 

The  niliilitiiin>  In  wliii-li  iiciimlir  liii'iithinu  nri-iiiN  inii\    lir  ili\iili'il  iiit" 

pli.\sii)|iij,Mi-    iiiiij    p,itli(iliii.'i''    i;liilips,      <>f    til,.    p|i\>i,iiiii,ri nilitiiiiis    tlir 

riil!iiwiii!i  nifiy   In-  tiikm  ;is  i-\;iiiipliN:   i  1  >    llri'iitliinu   in   :in   iitmiiNpliiMi' 
'•niifiiiiiinu  ii  <lflii"i<'iii-y   «(  <•  :   tlins.   piTimiii-  Iniiitliiiitr   is   viiy    n-iulily 
|hiii|iii-imI  in  p«'i-N(iii>  iivini:  in  rin.fii"l  ;iit         '     Tlii'  initinl  lircntliini.'  ml 
I'luiniriiii  ai)ii<'fi  iinlniTil  liy  tcni-iil  \  iiililat  imi  nl'  tlir  liiiiir>.     In  tliis  post 
.ipiicii"  |)i'iiiii!ii'it\-,  iIh'  apiii'ic  pi'iiiids  may  at    liist   1m'  ijiiiti'  niafkcd.  Iml 

as  lifcatliiiiir  ri'tiinis  lliry   li nii'   !.'i!iilnall\    slnntrr  ami    tlii'   liifattiiii',' 

intci-\als  iirailnally  Iihiuit.  nnti!  niMiiial  irspiratiun  is  ii'simi'il  i  l^'iy 
l-'Ii.  i:ii  HiTatliinir  tliiiiiii.'li  a  Imm  liiln'  liaNin^  a  small  \  csscl  I'lmtaiii 
!ii!r  Mil  la  liiiii'  insi'itril  Im'Ihitii  tin'  tuln'  ami  tlir  imiutli,  tin'  w  Imlf  i-apai-itx 
"t"  tills  \osscl  anil  tnliintr  licinir  aliniit  a  lit.T.  Tliis  «il|  caiisi'  piMtmlir 
'iii'atliiiiLr  in  piTsiins  tliat  an'  siisrcptililf  to  n\\i;cii  di'licifiii'v .  1C\  rn 
'•fi'atliina  tliriiiiuli  tlii'  tnlii'  witluiut  soda  limr  will  somrtiiiii's  cansi'  a 
iM'rinilii'  typi'  of  Ini  iitliinu  in  sui-li  iiidiv  idnals. 

Till'  imlliithif/i,  I'oiiditioiis  in  \v  liii-li  pi'iimlii'  In-i'atliiiiu'  lii'i'oiiii's  di'xrl 
■ipi'd  iiri-  ])aitirillaily  tlii>si.  assiM-iali'd  with  ii'iial  disrate  and  i-ii-i'lna' 
lii'imiffliairi'.      In    maiiv    ol'  tlii'sc  rasrs,    tlii'    pi'iiodic    Inoatliiii^'   diifs    not 

ipprai-  In  ili'prnd   on   tlii'  saiiii'    lai-lois  as  an nri'i-ni'd    '...    i!ir  i'.;-.    i- 

iiii'iital  txpi's.     Till'  sMiiptoiiis  would   latlii'i-  appear  to  df|)ind   mi       '  .i' 

iiiiilii'lli'i'  (li  tlii'  liiiiiiri    i-i'liliial   I  siipi  annrt>-a|- >   i-i'lilci-s  on   llii'   IrspiratufV' 

'•(■iiliT.     At  Irast  soiiii'  utlii'i-  i\  idi'ii t'  distiiflianri'  ol'  tin-  ri'ii'liral  fiiiic 

lions  is  always    I'm-tl mini;,   sindi   as  a    slii:lit    paialytii-   slioi-U.   and    tlie 

.'!71 


I'ln    i;i  -i-ii;  \ti(i\ 


piiiiMli.-    lllC-.llll 


iiii;    IS    iii-.iiU    ;il\\:i\' 


IIm'-c   cjims   ;i|v    Mlv,itl,\     l.rlic(it,.(l    l,\     ;i,l 


lUVl  ;iv  .lied    iliiiiiiy    slcc|i.      Mjin 


111  licilh   the  |iliysi..l.ii^i,-  :iii,| 


iiiiiKlr.-iti f  c.-itTfii 


'l''\''l'i|>    ,1    pi'iiodjc    cIlillMcIri-    i.iiK     w  li 


till-    Pilll|ll|(i;ri|.    n|(il||)s.     the    l)lc;itl 


miLT  iiKi\ 


I'll  till'   iii'isiin   is  .-isl 


I'i'li,  iind   cv.'i 


llnl|ii;ll     11 


|ii"|ili'.   |iillt|i-lll;ii  l\     mriilil^ 


•I'll.-ilh   (|i-'jl( 


i>    iilil    iM'oiilc,  in;i\    (Alilliil    it    I, 


h 


t:  ■  i    :  >.  i 


'M 


Causes  of  Periodic  Breathing- 

'"■'■'"      ""'■'■''^'     .-ill.i-llrs     l„     ,■11,      ll,V,.sti...,|li f     thl.     ,.;n|S,.S     „f     |„.,'IOlli.- 

'"'■•"'""-•  '""   ''  '■■'"  ' '•  ■■liii.M.'.l  th.-it  niiv    |M.i|-..,-tlv  satisr,-,i-t.n-y  IN 

'•'■"'^"'""''"^^'^>'''  ' n  ..llVii.,1.     IVnilni'.v  '  .-ilt  lilnit.'s'it  t„  ,,  .inish,.,! 

I'M'ii.iliililv   '••,   i.iis,.,!  thrrs |,   or  tl„.   i-,.s|,ir.-,ti,ry  iviifr  ,|.,..  to   fm.ltv 

l'l'""l     Mipilly,     .|„.     sU|,|M,sitin, ili^,     ,1,.,,.      „i,..„      ,|,,|s     s,||         ,.ss,.«l,      Ih'r 

""'""'•■'  ''"   "''  ''"■   '''"'"i    I-   'liLiiili'  to  ..x.-il,.  Ill,,  rcntrr,  so   tli        )„■,.'    mi'J 

Mops.     Diir.iiu  III,.  ,-,.sMl;ii,!.r  ,.||,||,.,.|.  en.  ;,j,i,i„  ;i,.,.,inniliit..s  until    ,,    li.is 


mil 


Tin:    IDSTIidl.    Ill'     III!,    lil.sl'lUAIlIlN 


i.iiscd  till'  ('ii  Mifliciciilly  to  CM-itc  the  (Icpics'-cil  (•(•iilci-.  ]  l>  |ifi])iicii 
inllows.  fiiiisiim  ,1  wiisliiiiy  iiiit  (pf  till'  Co.  .-iiiil  ii  rcMiltiii-jT  iliiiiiinitii)ii  i.i' 
ilii'  ctTcctivf  stiiiiuhis.  Ml  111   '  Mu.iiii  tlic  I'fiitiT  I'iiils  li.  Ill-  ^tiimiliilt'il  ami 

.■iii'|,.!rl  liii- this  i'\|t|jinali<iii  woiilil  a|)|>i'ar 


.•i|iiu'a  >U|>t'i-\  riii's.  ami 
III  ln'  I'liriiislii'il  li\  till' 
iiiLT  an-  iiiailc  to  liri'at 
(11,   tlic   pciiiiijicil y    111 


■t  ihal.  \\h.  !i   I     tii'iits  i'\liil)itii)_'  iii'i'iiiilir  lirralii- 
.1       iti^'iispl I'diilaiiiinu  ii   liiyli   pi'icciitaui'  nt' 


i.!i',-i;! 


iMiiii'.'  iiia\' 


liivc  iilari'  111  tfi,'iilai'  liii'alli 


a    ri'Milt    Nvliii'li    may    aJMi    .ir      litaiiit'd    li\'    iiiakiinr    Mir 


Ii    |)atu'iii> 


uniii'i-  I'asi'.  ilii'  si  iiiiiiliis  j> 


lii'ratlii'  in  atiiiii'>|ilii'ri's  rirli  in  iixyjri'ii.     in  tlii'  1' 

laiM'il  til  iiii'i'l   till'  (|r|ii  I'sscil  i'\i-italiilil\    of  tlii'  I'l'iiliT;  in  ihr  lattrr,  tlic 

(■\i-italiilit\     111'    llli'    I'l'lltir    is    ilii'i-rasi'il    lin-ailM'    nf    lirttcr    lilnnil    sU|i|i|\ 


tlial    it    is    I'liali 


til    irart    til    till'   iliiiiiiiislirij    stinmhis.      I 


III      I'M'll 


■-'lanti'il  tliat   tin'  rxi-italiilil  \    nf  tlic  i-ciitci-  is  ilcpi-csscil.   it    is  ijil'lii'iili    ti 


llV    tllis    sh.llllil 


siiin  a   pi'iiiiilii-  type  i,{  liicatliiiiL'  iiiili 


clinic  that    il    is  miU    w  lien  siimnliis  ■■  i. 


'•  .  <■,,  ,,r  III I      anil   tliirsliulil  i.r 


■  Aritaliility  111"  till nti'i-  arc  ailjiislcil  at  a  certain  pli>  sinlic^ic  lc\c|  tlial 

siiMinlli  ami  Clint  iiiiiiiiis  actiun  can   'jn  mi. 

Ilalilaiic  ami  liis  si-lmnl  a\cr  that    llicic  is  im  pcriiiaiicnt    altirat 


raliiiii    III 

'lie  cNcilaliility  111'  tec  center,  lr,it  that  tlic  pcriiiilicity  is  ihic  in  sc\eral 
causes,  uliii'h  ,1,1  not  alwa.vs  opciatc  to  1  he  same  ilcjice  in  tl,,'  iliiVcrcnl 
"■oinlitiiilis  ill  wjiich  sucli  peiioilii'ity  exists.  To  slllilv  these  causes  tin- 
c\;i,'l  I'omlitioiis  cxistiii'j  ill  the  various  tvpcs  ol'  perioilic  lircathiii'j  that 
can  lie  iirmlnc'il  cxperimenlallv    in  man  lia\e  liccii  iii\  estieatcil, 

'I'll,'    most    siiiipl,'    I sjili'i-    first    is    the    perm, lie    lircathinu'    that    i^ 

!'i'"'liic,',l   in  .-I  |.cis,,n  siisi'i'plilil,-  1,1  11     uant,  liy  liivalhine  ihromjh  a  tiilic 

■I'i'l    '"'"I I'    .-I     total     capacity     of     1     liter   ,     containing'    smja     rinic 

In   Mich   a   ,-asc  m,  oiitsiilc  air  enters  llit'   liiiiu.s,   I'm-   uliat    u  e  haxc   really 
Inne,  licsiilcs  pioxiiline    for  the  alisorplion  of  CO,,  is  trreath    to  prolon'.' 
•h,'  ,1,'ail    space,      'j'lic   ox.\  ^cii    teiision    of   the    rcliicatheil    air,    thcrel'.irc, 
'I'lickly  I'alK.  niilil  a1   la^t  a  point  is  rcachcil  at   \\liicli  the  respirator\    ecu 
ler  is  ilircctly    stiiiiiiialcl    li\    (I     ileprivat ion,   ;is    vvc    have   s,.cn    ii    i,i    l,,' 

'*ll''ll  this  falls  l.i  a  sliflii'lclillv  \,i\\  level  .see  pai;,'  :;.")n  .  The  ,|,','p 
meatlis  i  li,\  perpm  ;p  Hliicli  follow,  lieiiiu  of  greater  \oluiiie  than  lOllll 
■■  c,  eaiiM'  oiitsiilc  air  to  lie  iiispircil  so  that  the  O.  wan!  is  maile  "11111! 
iml  the  h.v  perpiiea  a'jain  ilis.ippears,  possililv  to  the  extent  of  alniea.  fol- 
low, in  ciiiiseipicnee  , 1 1'  a  coinciijcnt  "vvashini:  out"  of  I'O.,  there  has 
■"•1  11  a  low  CI  Ml',;  of  I  he  (  ',;  of  the  liloo.l  liclow  the  llircsholil  value  Dnrin.j 
'I"'  iipiica  the  o.  is  lapiill.v  use, I  up,  till  a  point  is  leachcii  at  which  ih,' 
•enter  aL'aiii    liccnim's   cxeiteij.      In    siidi   an    e.xperiinenl    the   clTcd    of  O 

'•'III'    I" im's    very    markcil,    as    show  11    liy    the    iiilciisc    cyainiMs    wliicll 

;.'.  i  Idiis. 


'I'liaf    lirealhiii'j    iiiiiler    t| 


coinlitions   sliiiuM    II,'    pciioilic    ami    not 


'#1 


lu'li 


:/ 


Till    i;i>i'ii;\THi\ 


1     f- 


mnvly  slinv^  ;,  >tc;i.|ily  iiici  .■;isil|.;  Iiy  i„.i|,nr;i  is  pruliilhly  <lll.'  I,,  tlic  ui 
•  ■.|U.t1  liitrs  ,it  ulii,.||  Ih,.  ()  ;i,|,l  (■(».  tensions  .•hiillir.'  ill  tlic  liloo.l.  I!, 
.•.•Ills,'  ,,r  ii  ••ImiIT.T  Jicti.iir-  the  hifl.i-  IliirlUillcs  iniK'll  less  tllllll  til,.  f,,i 
""■'•■       ■\ll"'l>il'   'MUs,.    I„,'   llir    |.,.rin,li,.ilv       is   l,.ul,t    Ih,.   ,l,.|i,y    ]H^t^^.■,■v 

tlic  -ii-  .■\clKiiii.r  in  th,.  Inii.^s  ;i!m1  til,,  .•iriiviil  ,,r  til,.  1,1 1  in 'til..  I,r;iii! 

When  the  (I    tensi,,n  m'  th,.  hi I  siipplx  ino  tli,'  n.s|)ir;|i,,ry  ,-,.|it,.r  fiill- 

'"  ^"  '""  'I  ''■^'■l  111'"  <'\i'it:itinn  ,,!■  the  e,.|,|er  n,.enfs.  th,.  ivsiiltiiiir  in 
'•n.iise,!  i.i-e.-ithinu-  iis|,ir,ites  ,,ntsi,|,.  (•    into  th..  ;il\,...li.     .\ft,.i-  m  iimiueir 

•"'  -"■  ''"■  ".    i-^  "■iirii,.,!   hy  th,.  M I   1.,  th,.  e,.nt,.r,  s,,  thjit    its  stiiiiiihi 

tion  h\  (I  ,hli,-i,.ney  is  iviii.,\e,|.  ini.l  it  is  h'tt  in  ii  ,.<ni,liti.>ii  in  \vhi..li 
If   I'iiils  tu  .jis.-hai-,.  iiny   inipiiis..^,  sin.','  thei-,.  is  .-i  snhnnnniil  C,,  „(  tile 

'''""•'  '■''■  "  '■•.iise,|ii,.n,- ■  til,.  l.iu.TiiiM'  ,,f  111,.  (■<»_.  i..nsi.,n  iif.Mln,.,.,!  |,\ 

til..    iiyi),.||.ii,';i.      A    lilt!.,    liiii,.    St    II, ,u    eiiips..    I„.f.n-..    lli,.    ( '(  I .    .-i^rjiji, 

iIm's  (II-  llie  (),   fiiils  snl1i.-i..ntiy   t.i  exeit..  th,.  e,.|it,.r. 


///,////L*//:i/##'^ 


a$.Mu.:M: 


illJUl  ,iJ' 


wwfwwvnfwn 


,t  '  •  ■ 


III  Ml 


Ull 


-      '  '■'       '-'^i.."li'.,lv.     111. ,1,1    ,1    1,,.   ,il.„i,j   ,,lr    tlir,.iiKll   .1   liil.r    _'(,ll   ,-,     l,,„„    •,,„!     ■   ci'i     ill   .Ir.iiul.  I 
■  lr,,:n    ll,.;,i;l:is    :„.,1    IlaLiUK.  I 

A  siiiiiiiir  Mlth.Mieh  i.'ss  iiijiik...!  ,|,..ji-,.e  ,,f  p,.,-i,„|i,.  i,f,.iitliin!.r  .•:iii 
sum. .times  l„.  .ihtiiin.'.l  hy  m,.ivly  ivspiiin.j  tiii-..iii;ii  ji  hmj-  Inii,.  withmii 
.•my  iii-..\isi.,n  fm-  tii..  .•il,s.,ipi  i,,ii  ,,f  (( »^.  hi  this  .-as.,  it  is  nimv  ,lirti,-iilt 
1.1  rxphiiu  111.    .■iiiis,'  ,,r  Ii,..  pei-i,„|i..  hiviitiiiny,  hut  that   th,.  xiiiin   Cai-t..! 

'""•■'■'"'■'I   '-  I'  *>    <l'piivati.Mi   is  .vi.i.n..,..!   hy   tli,.   fael    that    in   Ihi. 

Ms  ill   lii..  pivvi.nis  .•\p,.riiii,.|ii,  the  p,.ii.„|ie  iiiitni-,.  .if  the  n.spirati.m   i- 

' "''li'iti'ly   .■haiiu...!   t,,  Ih..   i..i:iiiar   lir..atliinu'-  if  ( >,  is  inti..„Iuc<.,l   int.. 

the  tiih...     Til,.  iiit,.|-,.st   ,,f  til..  .•xp,.iim.,|it    li,.s  in  liie  fact   that  a  siiniiai- 

n.iativ..   .■h.iiLMtein    nf   tl,,.  ,|,.,,|,|    spi is    pr.iiialiiy   a.'.-.iiintalih'   U<v   th. 

p..ii."lii.  Iii-.atiiin.^  s,.,.n  m  tii.'  \vint.r  sl,.,p  ,,f  tiii„.rnatiiij.-  aniniais.     Dur 

ill'-'  tiiis  e,„..|iti.,n,   <m   ,• ini   ..f  th,.  .|..pr,.ssi„n   .if   lali.ilism   |,.ss  (> 

'^    i''|i"i-,.|    ami    l.ss   (■(.     i.    pni.jn,.,,,!,    s.i    that    tl \eliaiii.-,.    ,if   ur„s,.. 

''"'""-''    ''"'    I'liliiimiary    .'mhit Ii,.|ii,m    is    ./ivally   .liminish..,!.      Tli.-   .I.'a.l 

^l'= ■    li"""'V<'i'.    i-ciiiains   .,|-   th,.    sain,,   .-apa-'ity,    « lih-h    ani.mnts    t.i    th.' 

^';'^'"   "■'—   •■-   if  ":•■   l;i"'''-  li;i(|    i..vn   i,,.,l.M,w...|    iin.t..,,   nii<.haii;:e,l   ,..iii- 
fliti.iiis  nf  pu!iiiiiii,ii-\    >^;\^  .'Ni'liaii",'. 


Tin:    (iiNIUol.   UK    llli:    Hl>l'lH.\TliiV 


'I'lic  cxpliiiijili.ii   lor  (illii'i    tyiM's  111'  I'xpi'riiiicntiil  pfiiiMlic  liicjitliiii 


ijrt's   occiiniii";   III 


iimcli   less  satist'jictiirv .      Inipoitiiiit   i'\  idciicc  tlint    dial 

till'   tcii.sioiis   (it'  (».   iiiiil   ('(',    ill    till'   iihi'iiliir   iiii-   iiiiil   tlu'rctoie   in    t)ii 


iiltTial  IiIihkI  oI  till'  iTspralitiy  ci'iitcr  ate  larjidy  ri'spiinMlilf  fni-  ]n'ri<Klic 

Hcathilltr  lias  I n  scciiiimI  1i\    stiidyiuy  the  cDiiditidi;  that  ilcvrlups  al'tcr 

I  iii'iidd  (It'  apnea  pnidiiccd  liy  \  (ihiiitai',\    lorccd  lucatliiiiii.     'I'iic  loMilts 
if  Miidi  (iliscrv  at  idiis  arc  u'ivcii  in  tliccur\ 


,  (■  >li(i\\  II  III 


I'isi.  l: 


'I'lic  tliiii   line  represents  tile  <K  teiisidii    ot'  tlie  al\('<ilar  air,  tlie  tii 


le   tlie  CO.   tension.      The   doulile   line    niniiiiiu' 


the  (diai'1    repn 


Milts  the  axcrayc  tension   of  ( '( »^   (iiirinu-  (|iiiet    noriiial    iM-eathine-.      The 
i(s])irat(iry    iiioveinents    are    represented    liy    the    traeinir    at    the    foot    of 


le  eiir\('  aldiiir  the  aliseissa.      It   wil 


lie   (ihserveil 


that    the  owncn   ten- 


sion falls  \cry  rapidly  during  the  apiieie  jieriod,  until  just  liet'ore  Iireath- 
iiijl  reeoiiiiiiences  it  may  lie  as  low  as  :'>()-:!.">  iiini.  liy:  instead  of  the  iior- 
iiial  (if  alioiit  !(.">.  .Meanwhile  the  Ctt.  tension  rises  from  the  very  low 
le\cl  of  1"J  iiini..  at  first  very  rapidly,  then  more  u'radiially,  altlioiifrh. 
when  lii'eatliiiiir  reconimenees,  it  has  not  yet  eaiiied  the  normal  level. 
.\s  a  result  of  the  first  perio(|s  of  lireatliiny,  the  (»,  tension  suddenly 
shoots  up,  lint  the  ('<>.  falls  oiil\  sliu-litly  During-  the  next  ainieie  staL'e 
the  ()_.  (piickly  eiinies  down  ayaiii,  and  the  •'(»,  rises  so  as  almost  to  ;*■ 
tain  iKirmal  tension  lief'ore  lireathinu'  airaiii  supervenes.  .\s  the  api. 
periods  sulise(pieiilly  lieeome  less  pronounced,  the  ('( • .  tension  comes  to 
stand  almost  at  its  normal  level,  whereas  coiisidcralile  variations  in  the 
<>,  tension  continue  to  occur. 

Several  interest iiiir  t'eatures  of  these  results  dcuiand  attcnti'iii.  In 
the  first  place,  it  is  plain  that  the  Imdy  is  jiosscssed  of  smiie  niccli.inism 
hy  which  it  can  ])re\('iit  irrcat  tliictuatioiis  in  the  ('(>.  tension  of  the 
lilood,  whereas  towards  (>.  no  sin-h  "liulTer  action"  is  displayed.  It  will 
fiirtlier  he  oliserveil  that  the  ('<  >^  tension  of  the  alveolar  air  rises  \  cry 
rapidly  durins:  the  first  part  of  the  a|ineie  i>eriod,  and  then  iiior(>  yrad- 
ually,  the  ex]ilaiiation  lieiiu!  that  durin-r  the  foiced  lucathinur  the  CO. 
has  licen  washed  out    from  the  lilood  hut    not    from  llie  liody  as  a  whole. 

.\t  first  siiilit  one  misrlit  attrilmte  the  periodicity  to  the  same  cause 
as  that  operating'  durini,'  hreathin;;  tlirou'_di  a  lonu'  tulie  w  itli  soda  lime 
namely,  to  oxyireii  deficiency.  Miit  this  explanation  is  unti'iiahle.  lie- 
caiisc  the  perio(Iicity  remains  evident  t'or  some  time  after  all  possiliility 
of  direct  stimulation  of  the  center  of  O.  ileficieney  is  over.  .\  possihle 
clue  is  furnished  liy  the  fact  that  lireatliiiiy  returns  while  the  CO.  ten 
sioii  is  still  consideralily  lithnr  its  normal  level.  The  vtiirii,  as  we  liavi' 
seen,  is  accounted  for  hy  the  app<'araiiec  of  hactic  acid,  and  if  wo  assume 
that  this  has  (icciirred  particularly  in  the  respiratory  center  itself,  a 
slitrht  deirree  of  hyperpnea   will   he  cxeite(l.   which   I'v    supplyiiiL.'  O.   v\ill 


iril^lil 


PL 

mm 


I 


i    I 


Ih'l- 


:i7(i 


■I'm:    KISIMKATKIV 


11' 


il  -"^ 


^      1 


»-! 


4-  ' 


i 


■|>i   '-Llv     nNMllZ,.    tllr    la.'l  ir    ari,|.    M,    ll,;il     Uu-    still    >|ii.|it|y     s,|l  .     .nilid     . 
"'    ""■    ''' '    '^    ""•'''!-•    I"    ''^'-it''    II"'    '•'■HIrr.        Al  llfi,    th.MTlnlT    .SU|»TN  m. 

••iM.I  hists  III, III   hicii..  ;i,.i,|  Ii;is  .-m.-iiii  iirrmiiul.ii,,]  ill  tlic  .•.■liter.     T..  .- 
I'iiiiii   "liy   l,„-;i|  .•i.-.Miiiiiihiti.m  n(  la.'lir  ,iri,|   in  tli..  .-.•nt-M-  slmuM  lirclu.- 

"   l"''i'"li''  'M f  I'lvaljiiiiu,   „,,   „||,^,    |-|,|.,|,,,,.  .|^^„ ,1,.,,    III,,,.,,   j,.   

siW.raM..  ,|,-lav    l.,.lu...,i    tli,.   iM..|.i.Mit    al    ulii.-l,   .•.|iiilil,riii„i    „f  tli..   ..ras, - 

'"    ""■    ''' '    ■""!    ■'I^''"l.-n-   .-111-    Im.,-,„ii,.s   rstal.lisi an. I    tiiat    at    uliin 

''"■    ''' '    ■"■'■i--'-    ■"    III''    i"s|.iratniy    ,-,.,, ,,.i-.      Tliis    ,|..|ay    is   .■allM.,l    \:^ 


til.'  nI,,\\  iii.j   ,,r  tl,,.   M hlnv,    uii   a, 


IllnV  ('|||.'llt> 


lint    III'  till'   ansi'ii.T   III'  f.'spiratc 


Kliililiasis   is   |,la,-,..l   ,,,,    th,.   tact    that    it    is   in   tli,.  ,M.|it,M.   its,. If  an, I    im: 

'"    ""■    '•' '    '!'■"    'I'"    l'"-'i''    ^"i,!    l„...,„ii,.s    ,,xi.li/..,l    i,y    til,.    ..N.-.'ss   .,1'  (. 

I'-'Mils,.   Ia,-ti,.  a,-i,|   is   kiMUM   1,1  .|isai,|„.;,r  s|,,„  |y   ni,,!..,-  tli,.s n-iitim.. 

"■ i-''.""i  i-1 1.  I'llt  t,>,l„s,,v,r.v   ,|ui,.kly  fi-niii  tiss,i..ss,i,.ji  asnius,'!. 

.'Ill'l     I'll   sill, ,;,i,|v      til,. I,, I',, I-,.    -iK,,    |',.,„||     II, .n,,,,..,    tissi,,. 

Ill       S|,|,|„,,.,       ,,1       111,,      .||„„,,      ,.^|,l;,|,;,,j,,||       i,       1,.,^      I,,,,, I,       |.,„|||,|       ,,|.|,  .|.      ,,,^^    .,,,,, 

""■  -'"'l   "I    'l"'    i"i'-''l   l.n.atliiiio    tl,,.   Iiino.   ar..   (ill,..l    „  JH,    suffi,.i,.„t    «i 

"'  '''■"   ""■  '""-i"ii  "I'  lM>  ^.'is  in  til,,  alv i  is  n,it   I.iu.t  tliaii   120  niii. 

Ik',   iMvatliin.^    is   i-,.;.r,i|ar   i„   ty,,,.   „1,.,,|   j,    ,,,,,|,ns,  aii,|   tin-   CO.  t..Msi,„: 

"'■  ""•  ^iIm'"1;ii'  ^lif  is  s,.v..ral   iiiilli,ii,.t..|s  „h„r.    iiist..a,|  .,t'  l.,.l..«'tlu,   

iii.il  stiiiiiiL-itinu   l,.\il. 

'•'"   -'""    "!'•   ""•    l"'i'i'"li''   ''liaiacl,.!'  „(  t||,.    I,|...atliiim    sii|,..,.vcniii.,'   ,n: 

•■'.  '"''■""'  '"■  "l""'^'   '"">    ' M'liii I  ^is  f..ll..us.-    m.l,.,.  .M-.linai-v I, 

'"""  '"    '"'■■"'iii'^   ^'ii'l    I'.'inMii.'ti'i,.    inrsMir,.   tli..   (»,   fiisi ,f  tix"  I.ln,,,! 

p  suit,,.,,,,,   l,„uw...M  ,,„,,„a|   ,,,  ,.in,ti„ns  t„  ,,f..v,.,it  anv  a.-.-,nnulati„n  n. 
I'i"'i'-  .'i-i'l  111  tl,,.  i-,s|,„;,t,My  ,-,.|it,.r.  s„  that  tli,.  stinuihis  alV-.r.!...!  i,v  ll,. 

'  "    "'    ''"'   ''' '    I"'"'' -   ■'    ''"iistant    ,.|r,.,-t.      Diiiii,.,.   th..   apn..a    Mlii.-h 

-U|...r>,,i„.s   ,i,„„i    ,„,v,.,|    ln...athinjr,    la,-ti,-   a,.i.|   a,.,.|iiiiulat,.s   in    th..   .-...it,.!- 

■•aiisi,,...  tliist,.  r,.s,,„ii,|  t„  III..  ura.iMallv    rising  ('„  „r  th,.  1,1 |  l,,,,-,,,,.  ,„, 

Iji"..-   has    ,.,,i,.h,.,|    its    |,|i.si„|,„i,.    l,.v,.|.      Th..    hy,M.|.pn....,    thus    ..x,.it,.,l 
•lo..s    i„„.    |„,vy..N..r.    I,nn-   ,ilM,i,t    ,■,    i,,-,,,,,,,)    ,,Ni,|;,ti,.ii    .,f   tli,.   la.-tic   a.'i.l 

'"  ""•  "■'■'"'■'■  '"■  ■•'  l""'''iii'^  "I-  Ilir  «■„  ,it'  th..  1,1 1  ,.i,...„|ati,m  tlii'.il|.'h  it 

"'■■""" '■'•   """■   '1'""    "^"■'1    i^   '.•il<''ii    lor   111,.   1,1, „„1    t„   J....    fr,„M   th,- 

'"'-'  *"  "'"  '"■"'' ' """I   '"■  'll"  .'il'-'ii f  r..spirMl,.rv  .M..v,.|n..nls 

^^'"■"  ""•  ^"■'■^"'"'  ''' I  '!"-  "■■■I'-li  111"  i'.'spirat„ry  ....nl,.,-.  fh..  ..x....ss  „. 

"■   "lii"l'   i<    -Mitains  „xi.|ix..s  ,h,.   la..li,.  a,-i.l   s„   that    ap„..a   s„p..|.v,.„..s. 

;""'    "'''   '"'•'"•   ■•"■"'   ■'-•''"    .•i""'iMiiilMt,.s.   altl L'h    ii„t    „„„    s,.    ,iiii.-li    .■,. 

■  l'"l'""  l'"".iiiM.  „1'  th,.  ..la.lualK    rising  ('„  „|'  th,.  I,l„„.|  its..|r.     Th..  ,.ss..n 

"•■'I   '""'"I-  111   III..  .■ai.sati.Mi  ,>t'  p,.ri,„li,-  l„.,.Mtliin-  is  tlM.r..|'.,r..  a  ih^lav..! 

mass    iMii\  i'i,|,.i>*    .|i'   .(,,.    1,1,,,.,!    I',.    ,      '1       1     I  ■>.      - 

....".■(    ,i,,i,,    :!;,.   j,alninii;ny   .-,, j.itlaii..s   I.,   ih(,   p 

s|iir^ilui  y    ...nt,.,       Th..  .l,.|ay  mav    I,..  ,.a„s,.,|   l.y  ....ssati,.,.  „|'  tiu-  r,.spira 


Illi:    CIINIIidl.    n|-     Tin:    ICIM'lliAIIIlN 


lory    lliovclilfni.   il-   ill    |ii]sl;i|)iiric    |>c|ii)(licity.   (, 

riiliitory  ilistiirli;iiii'i'. 


1-   \i\    xiiiic   |iiiliiip|(ruM('   rir- 


l'ci-i(»lic  lircitlii 


ii'j-  IS  |)i(j(|iii Iiy   riMi-cd   ics|)iiiitic)ii  inufc  icadil 


l.lli'fic.l    ilil-  tll,'lll    111    sc;i    lr\..|.       It    v^iis    101111,1    l,y    1  )oiio  li|.,,-'    jifl.T  l)rciltl 


III".'    Iid'cililv    for   III 


ic   limiiitc   ;il    sf.-i    Icvi'l,    tlint    Ilic    I 


icliiriicd  sliowtil  s  1,1   10  ,ij|-|' 


MCMlllllli;     wllfU     it 


'\|>('niii,'iit    ill    iiii   ;illilii,|,'   iriviii>_'   ;i    luir,, metric    p 


ll'|l,'tltH)ll    ,il'    tl 

Miiv  of  (iOd  niiii.,  lTi  Mu-h   |(crio,K   f,i!lou.>,l   t 

li'spl 


Vri'ht   |),Mio,ls  (if  jipiicji  Jill, I  liypci  piicii.     () 


II 


iiiilinu:  to   ")2(l 


iiiiii..  4(1  p('ri,>,N.      IihIi'cI.  jit    liiul 


H'  ;ipii,'a  :   ill    M    licmlit   cir- 


1   ;illitii,|cs  pcii,),li,- 


iifiitliiiiii    iiiii.x    \n-   lin.iiLdit    iili,iii1    \,\    III,,   sii•illt,■^l    ;ill,T:it 


ion    ill    iioriii.'il 


JltM.II  ;   v\ru    tuUill^r  ;|    ,|,.,,|,    |,r,.;,th    lllliv     ll,.   Mlfli,-i,.|lt    I,,   .Mils,,   llistin,'! 


Ill    iiMiiy    pfrs,)iis   !i\ 


p,'rii),|icily    III   111,.   sii,.iT(.,liiiM    i.,.>piriiii,,|is.   jm,! 

nt   liiu'li   iiltilii,|,.s  p,.rj,„li,-   i,i-,.,.|tliii,._r  is  very  ;ipt    1, ,.,ir  ilmiiiir  ^Ic.p 

As  ill  piilli,.|,,.;i,-  ,.;is,.s  ,.Nliil,itiiin  (■|i,.yM,.-St,,k,.s  r,.spir;iti,,ii,  tin.  p,.ri- 
o,li,-  l.r.'Mlliiii-  ;il  hi-li  .•iltit,i,|,.v  ,.,.,11  |„.  iiiiiii,.,ji,.|t,.|\  r,.|ii,.v,.,|  l,y  iiispir- 
ill^r  iixy^.n, 

W'l'     ilil\(.    ,|,.\Ot,.,l    ,.,l||si.|,.|.,||,|,.    spil,.,.     1,,    ;,     ,|iv,.|ls,si, f    ll|,.s,.    ,.Mr,.„„.l\ 

■lil'lil'UJt      prol,l,.|||S     ill     111,.     ||,,p,.     I||;|I     ,.|i||„.;||     ,,l,s,.r\,.rv,     i.y     l„.,.oiiiiinr     ;|C- 

'I'i'li'i'''''    "I'll    '111'    |iur,.l>    t.\p,.riiii,.|il,ii    u,,rk.   iii;i\     li,.    in   ;i    p,,siti,,ii    I,. 

' '"'•'    'll"l<'   s,.iir,-liil|o    ili\,stiii;itiolis   ;|s    I,,   til,.   ,.,iu>,.   ,if   ( 'Iny II,.  Si ,.k,.s 

.•mil  iitlicr  p;itlii(loyii.   forms  ,if  p,.ri,Mli,.  lir,.;it  liiii'.'. 


:.-if 


i  f 


Is 


1     'il 
1     t! 


I'Mj 


Sir  - 

f  ■If  I 


v-\  a 


I:' 


rllAI'TKi;  XI, IV 
RKsl'IU'ATKtN   |{|;Y(»\I)  TIIK   MN(.S 

I  p  III  thr  fircM'iit  oiii'  sill. lies  ill  r.s|iiriiti(iii  have  ((iiii-irriiMl  the  varinii. 
;ii(<'liiiniMii.s  iuviilv.MJ  ill  liriiijfiiii:  almiit  a  ciiiiNtant  (•liaii<;f  in  the  nun 
iw.sifi.iii  (.)■  ili(.  alveolar  air.  \\\-  nnist  now  consider  the  nature  ot'  tin 
-.iieans  \n  wliicli  ilie  ox\  ireii  is  eoiiveyed  to  the  tissues  anil  tile  CO.  re 
nioved  t'riini   tlieni. 

In  the  first    plaee.  it   is  iiiipoitaiit   to  note  that    it    is  not   for  purpose. 

"'■  "Ni'l.'itioM  in  till'  III 1  itself  that  the  ( >,  is  reipiiri-d.     In  its  respirator.x 

fiinetioii   this   (liiiil   serves  as  a   transiiortinu;  aireiiev   lietween   the   lun-- 
and    the    tissues,    in    uliidi    reside    the    fuiiiaees    of    the    liody    that    eon 
suiiie  the  O,  and  produce  the  (■(»_,.     This  does  not    imply  that  tliere  is  im. 
oxidation  ill  the  hlood  itself:  indee.I,  we  slioiil.l  expect  a  eertain  deuTe 

of   oxidation    lieeause    of   the    fact    that    the    hi I    contains   s e    livin- 

rells  the  leucocytes.  It  is  scarcely  necessary  nowadays  to  offer  e\  i 
deuce  for  the  forcKoin!:  conclusion.  ( tne  well-known  e\|)eriiiiental  |)iooi 
consists  in  n  iiloiiii,/  th,  hlnrxl  in  ,i  fn,<i  with  iilnisloliif/ir  sullii,  soliilior 
iind  then  siil,.jectin!j  the  fro-:  uilh  the  saline  in  its  hlood  vessels  to  ar 
atiiiosphere  of  pure  (l„  when  it  will  he  found  that  the  animal  continues 
to  alisorh  the  normal  amount  of  (»  and  exiiale  the  norinal  amount  oi 
<(>..      It   lespiies  iiniiiially  without  any  IiIo.nI  in  the  liloml  vessels. 

In  order  that  this  transportation  of  L'a.ses  lietween  the  luntrs  and  th' 
tissues  may  he  .'fiiciently  pcrfornied,  the  Mood  iiinst  he  |irovi<led  will 
means  for  carryiiii;  adeipiate  amounts  of  ^'ascs  to  supjily  the  reipiirement-^ 
of  the  tissues.  Imth  durinir  rest  ami  during  their  varyin;;  dejrrees  o! 
activity.  .\ot  oiil.v.  iheret'ore,  must  the  (i.  and  ( '( l_,  cajiaeity  of  tie 
hlood  he  very  eoiisiderahle.  hut  it  must  he  capalile  of  very  rai>id  a.l.jusi 
nient  from  time  to  time. 

••iir    prohleiii    naliirall.\     resolves    its,. If    into    lliree    .larts:    iH    the    cal 
of  the  tissues   tor  oNXMrii    I  lialvrofl  I  :  or.  as   it    is  st.vle.l.   tissue  or  inlernn! 
respiration:  ' 'J     the  mechaiiisiii  hy  uliidi  the  hlood  transports  the  prope 
Jiiiioiiiits  of  ..-ases  to  meet    the  rei|uirenients  of  the   tissues:   and   CO    tli' 


nici-haiiism    In     whieli   the   Mood 


tel  Hat 


nil   respiialioii.     For  conv  enience,  however. 


rises  are   exchaiiL'ed    in    the    lun<rs 


ur.il  onliM-  and   consider  the  transportation  of  tl 


we  shall  chanire  this  nat 


le  irascs  lirst. 


UI.MMH  \ll(l\     lil  \llMl     llll,    I.I   \(is 


:n9 


THE  TRANSPORTATION  OF  OASES  BY  THE  BLOOD 

The  Transportation  of  Oxygen 

It  is  |)l;iiiily  lint   liy  iiicic  Miliitiiiii  in  tlic  pliiMiiii  i.r  the  IiIi.chI  tli;i;  tlic 

(liiiispiiitjiti r  O,  (H-ciiis.   I'lir  iit   the  itiiitiiil   i>i-cssui<'  nf  il.is  >,'ns  .'X- 

isliii!,'  ill  tlif  jil\     Jjir  iiir  at  tlic  tcinpcriitnrc  of  tlic  ImhIv  the  aiiniiiiit  lliat 

iiiiild  l)c  (li>s(.i\»(|  ill  tlic  III I  woiiM  he  (Hily  ipiic-fiiiticth  (if  that  which 

In  actiiallv  I'diMid  to  he  |)i-(scii(.     If  there  were  oiil.v    plasma   in  the  Idixxl 

vessels,  it  wonhl    require  a    viijiinii.  uf  lliiiij  a untiiiM   Id    l.'.ll  kihiu'ianis 

iir  more  in  order  to  convey  the  necessary  aiiioinit  of  (>     fnim  the  liincrs 
lo   the   tissues;   that    is,   tiic   coliteiits   of   the    vascular   system    uolllii    wci-;!! 

tvviec  as  much  as  the  avcraire  wciulit   of  a   man. 

The  suhstance  that  carries  the  <  >^  in  the  lilood  is  the  In  nKnilohlii.  \vlii(di 
may  he  dcscrihed  as  a  hiirhly  complex  iron  <-ompound  of  protein  espe- 
cially evolved  for  the  piirjiose  of  transijortini,'  (V.  In  some  of  the  lower 
animals  other  compounds  exist  in  the  lilood  f(M'  this  |)uiposc,  hut  none 
nf  them  is  to  he  compared  in  its  enticieiicy  with  heiiioMlol.in.  They  are 
merely  jmor  imitations  of  il. 

Uevrardiiii.'  the  conditions  under  \vliich  lieiiionl,,hin  comliines  with  or 
delivers  up  (>,,  the  first  (|iicstioii  that  presents  itself  is  whether  or  not 
the  reaction  is  a  strictly  (dicmical  one,  If  so,  a  dclinitc  amount  of  O 
must  he  capahic  of  comhiniiij;  w  ith  a  delinite  amount  of  heiiio>rloliin.  It 
Is  impossihie  to  secure  licmouloliin  of  siifticieiit  purity  to  test  this  rela- 
noiishii)  directly  on  hemoirlohin  itself,  s.i  that  we  must  iest  it  indirectly 
liy  examiniiiir  the  comhininir  e(piivalcnt  hetwecn  (),  and  that  i)oition  of 
tile  hemofilohin  molecule  upon  which  the  comhinint;  jxiwcr  deiicnds.  This 
is  the  ]iart  of  the  molecule  containini;  ii'  n  N'ow .  if  we  com])arc  the 
iiiioiint  of  (),.  \vhi(di  hemotrlohin  can  taUc  up  with  the  amount  of  iron 
present  in  the  hcnio<;lohiii,  we  .shall  find  tiiat  nui  nloni  of  inm  hniiini.^ 
ciimh'nifd  wllli  two  aloiiis  of  O,.  Evidently,  then,  we  an-  here  ilealin"; 
■Hfh  a  dctinito  chemical  reaction  occiirrinir  hctueeii  the  O.  and  the  iron 
of  the  hcmatin  portion  of  the  hemoudohin.  This  ndationship  is  known 
as  "the  specific  oxy<:en  capacity  of  hemoirlohin.  " 

In  showing:  that  the  union  of  (».  and  hemo<rloliiii  occurs  accordinu'  to 
'liemieal  laws,  ,ve  throw    into  promiiicnee  consideration  of  the  mechanism 

hy  whi.di  the  (>,    comhined   with  lii'moulohin  in  the  lil I   i.s  raiiidly  dc 

'ivcred  up  ill  the  <'apillaries  so  as  to  sujiply  the  tissues  w  ith  their  rer|uire- 
iMcnt,  and  is  then  as  rapidly  rccomliincd  .,i;ain  in  the  lun«s.     Moreover, 


'c  must  reconcile  facts  implied  liy  the  i<!ca  of 
lie   well-knoun    ohservation    that    tin 


II   sp. 


•itic  <  >_.  cHpaeity  with 


le    iiemoirlohin    in    the    circi 


dat 


ion    IS 


isiiallv  united  will 


1  coiisidcrahly  less  < » ,  than  the  total  amount   po.ssihle 


i 


I 


1   'j'l 


:!M) 


nil     Ifl.M'lUATloV 


ii-^^! 


I? 


It'- 


MM 


■^f  <' 


M 


In    (piIht    wor.ls.    VM.    i,iii>l    irc.-ni/,.    ih.-il,   nlllMMiyli    it    is   fssnif ijill.v 
■li-lninil   iciirli,,!!.  th.'  .'(.iiihiiiati.Mi  ufO,   with   li.-im.i;l.)l,iii   is  j;i-,.;itly   ii; 
llii.Miccd   l.y  oili.r  iMi'lois.  iiiHl   Hint   it   is  tli.'sc  tluil   aiv  liUoly  t.,   '„•  ,,: 
|)li.\  si(p|cii.'ii'  iiii|iiirtiincc. 

Ill    Mi-.l.T    tn    lllMlcisliiiMl    /,.r    ,„,„lili„„s    Hii.hr    ,rl,irl,    In  niofilohln     ,r.r 
t<ik,    I,,.  ,nu/  .,ir<    oil   O     in   lln    nnnn,,!  h,„hf,  «  ,.  ,„„st   sUl.jy  ill,.  .M.iiihiliih- 

'"'""■'■  "'■  •"■ i-''"''i'i   "li''"   It    is  cNpos,..!   to  .iiirnnit    iMilial    pirssMi, 

'"'"-■  "■'"•  I'l"-  ;.">v"nmin-  this.    s,...    ,,,„;,.    :::;,;.     i„    ,|„-    i,| |.    ,|„,   , , 

fivnirs  of  the  partial   pivssiur  <,(  O,  aiv  tvpirsciitr,!,  at   Ih, ,.n,I.   I, 

that  in  \hv  ah,',.lar  air.  u  hi.'h  « .■  haw  -nn  lo  !.,■  ahniit  jllO  mum  II- 
'Mi.l  at  the  other.  ]n  that  ..sistin-  in  th,.  tissues,  s.i.-h  as  ninsH.'.  MJii.-i 
has  !„.,., I  shnuM  i„  |„.  „ot  inoiv  than  l!l  or  -Jd  mm.  Ifir.  We  must  fiirtli, 
l-.-n-  in  "liinl  that  tii,.  n,  i,,  its  passage  from  Ih,.  alveolar  air  to  th,-  Immi,.. 
irlohin  an,!  fr,.m  th,.  h,m.o-l,.l,in  to  tli,'  tissm^s.  is  transmit^,!  in  soluti.. 
1hr,.ii!:h  the  jiiasma;  that  is.  so  lar  as  tli,.  supply  of  (».  to  tlie  tissue  rv\<- 
is  .•oiie..riHM|,  th..  plasma  s,.r\  .'s  as  th,.  imm,.,lial,.  s,.ur,v.     Sin,-,,  th,.  ti^ 

su,.s  ar,.  usini:  up  n.  ;,!  ,.,  v..ry  -r,.at  sp,.,..|,  ,.sp,.eially  «h,.n  aetiv,.,  , 

iii-e  thus  ten, linn  t,,  1,,^,.,.  ,1,,.  t,.Msiou  of  <»,  in  the  plasnm.  fav,.rai,l,.' ,.on 
-litioiis  hav,.  to  Im.  ,.r..al,.,|  u  lu'rehy  th,.  h,.moL'h.l.iii  lil.,.rat,.s  ().  at  tl,. 
Nam,,  rat.,  as  that  at  uhieh  it  is  h-aviii-  tli,.  j.lasma.  In  l.rief,  i"t  is  tl, 
O  t,ns„n,  „f  th,  i,h,s,n,i  in  Ih,  l„„„  ,„,,iU„ri, .  Ih„l  is  Ih,  i,„,„,rla.' 
f'lrh.r.  Ih..  li,.nN.M|„|,iM  „i..r,.ly  s,.rvin..'  as  a  stor,.h,Mis,..  uhi,.),  ,|,.|iv..,~ 
lis  <>,  at  .just  su.-h  a  rat.,  as  t.,  inainlalM  th..  plasma-..xv -,•„  fusion  a! 
"  •••"'•^f'""  l''^''!  It  i-  ol.\i,.us|y  of  th..  .,'r,.at,.st  imp,.r(aiH-e  that  av  . 
sh.M.hl  nn.l..r.stan,l  h..«  this  m,.,.hanism  of  an  a,|,.,p,ate  plasnin-oxvii..' 
ti.iision   is  iiiaiutaiu..(l. 

Methods  of  Investigation.      WV   must    nmemher   that   the   ,-,-ml.inati,., 
ot    <•,   an,l    h,.mo-l,.liin,   l„.ini.    a    ,|,.|init..   eln'mieal    r.'aetiou.   will    l„.    i, 

^.•|■siMe.  ami   must.   th,.r,.tore,   ol„.y   th..   laws  of  mass  aeti s..e   p.p.. 

-'■■!i    a..,-.,r,lin-  to  th..  ...piath-n:    III,  0,^I1I,(),.      hr   ..r.l.r  to  as,.,.rt;,n 

th,.  p,.sition  .,f  th..  halan..e  of  this  ,.,, nation  at  ,litrorent  i>artial  pr,.ssnr. 
ot  O.  that  is.  th..  r..lative  .piautiti,.s  ,,f  ,,xy.  an, I  r,.,lu<',.,l  h,.iii,.-.'l,,l,i. 
1orme,l  iu  a  s,,lution  ,>f  h,.m,.i.'lol.in  «  lu'ii  this  is  shak.-n  with  (».  at  ,litV..i 
ent  l)r..ssur,.s.  u,.  „uiy  pro..,.,.,|  as  to  h.ws:  A  lew  e.e.  of  the  heii„)!rrl,.lii. 
N..lii1i.m  ar..  pla,....l  iu  .aeh  of  a  s,.ries  of  v,.ssels  eall,',|  I,,,,,,,,,.!,,-".,  lik. 
those  sh,.«n  in  Ki.^.  i:!4.  |„  a.j.lition  to  th,.  h..mo.j|ol,iM  s.ilulion.  ....i..! 
t.mom..t..r  .-..nitains  a  mixture  ..f  nitro-en  an. I  (»_.  j,,  ,li|T,.r,.ut  iu-op..i 
tions,  Supp.,s,.  u..  us,.  MX  v,.ss,.|s  an,!  iu  .\,..  I  have  pur,.  nilro..M.|i;  n 
No.  ■-*,  nilro-;,.u  .•onlainin-,'  .")  mm.  parti.-i!  picssuiv  .if  (I  ;  in  .\...  ;!,  |.^ 
mm.;  in  \o  4,  -Jll;  in  .\o  .-..  .-,();  ;,u.l  in  .\,r  ti.  jlio.  We  nou  n.lale  tl,. 
Ionom,.|,.rs  in  a  uater-hath  at  l.o.ly  t..mperature  for  al.oi.t  lw,.nty  u,ip 
iil.'v.  s,.  that.  \n   th..  lormali.ui  .,f  a  tliin  film  ..f  h..iiio>r!o|,i„  soliithm  ..\.- 


.•I  ^I'li;  V  Tins    i:i  \.iMi    INI     I  I  \i; 


:!Ml 


Mil'    UJllls   III'   llic    VCNM'I,    pi'llVi-I    ri|l|ililj|  jiiiii    li 


lliiiM  iiiiiy  III'  Mitiii I 

liill    snlllliiili    lit]     III'    III 


I't  W  I'CII    llic    jlllllllsl>ll('ll'    ,'ll|l| 

A   iiM'ii^iil'i'il  i|ilinitity  III'  lii'iMii- 
<•  ('  1    is   tlii'ii    t'l'iiHiv  I'll    1111111    I'Mcli    liiiiiiiiii'ii  r 


(  si'i-  ]>;]"{•  :;.iS  1 , 


I      ■:'■ 


Vi'.'i;    l.v     I.  |.  .,tl.,l,i 


.  inl    |il;ii'i'il,   li.ycllii'i-   \,itli   miiiw    Vi'iy    ililiU,-   imii i:i    In    hike    llic    li| j 

II  iiiit'  III'  tlic  siiiiill  lintllcs  111'  ,lu'  iliil,  n  iilliil  iiKiiioiiii  h  r,  sliiiwii  in  l-'i^. 
i  1">,'  'riiJN  iiiiiiiiiiiicfci-  I'liiisists  ill  ])riiii'iplt'  ii|'  m  LMiiiliijitcil  I'  sIimjh'iI 
•iilir  III'  iijiniiw    liiiic.  I'liiitjiiniiii:  i-lovi'  nil.   tlio   I'li'i'  iinl   ,,(  \\,r   I'-tulii' 


l.'v      li^ii.  rcill  .    lii'l,  I-,  iili.il    I.:. .. 


I     u.i>     M);inni 
■I    I., .Ml.  .    -h. 


•  I'l         11  <■    ..u.ill.i 
'.I.I  I.,-  .l.-.,.,r.'     r. 


f  l;,I.,-    ,-, .,.':„„-    ,|,.u     ,„I. 


■in<;  coiMioctcd  wilii   siiiiill   lidtflcs  inoviilcil   with   snnic  d 


fluids 


('Vice  so  that 


111   he   placed   in   each   nl'  them   Imt    kciil    iiimiixed    iiiitil   tl 


'ttle  is  viiileiitlv  shaken.     The  tl 


IP 


iree-\\a\    stiipcock 


liet\ 


the   small 


1       I 


riu-  hi.., I.) 

•I.  rtwiuMtivrlv.      T 


K;i-    m.itinmitcT -i   art     umiI 
hv  r<-iiilts  Willi  ill. 


-iii;il!  Mii.iiililii 


I'll     iiM     uiih     I     r  .  .    .-111.! 
aic  .1-  ;ii  I  ii'  :ni    .1-   «  i 


TIM     HI  ^ni;  \rln\ 


I;     •■• 


IkiIIIi'v      ;iII<I       IIk'      lll.'lllnllirll   I'      s('|'\  I'N      III       |II'|M||I       I'llllllllllllil-ilt  inn       III       ill! 

IIIMMiilili'trl    vv  il  ll   tl llNiili'  ililV 

All    rijiiiil    i|iiiiiitit,\    III'   lii'iiinuliiliiii   Miliitiiiii    lliiit    hiis   I II    kiitinjiti''! 

Willi  ii\\l;cii  i.e.  ii\yliciiiiii.'liiliili  i>  |il,irii|  in  llir  Imtlli'  mi  llii'  utln! 
■  'III!  Ill'  tin*  liiiiiiiiiiii'tiT  liilii'  ri'iuii  tluil  ciiiiliiiiiiiiL:  IIk'  I'litlli'  uitli  tin*  iiii 
s;itiiiiit<il  liciiinirlnliin  Miliiliiiii.  'I'll'  liiittliN  liiiviiii;  Ikm-ii  jittiiclu'il  I. 
llir    iiiiiiiiiiiictir   \(illi    till'   sliipi'iirks  ii|n'ti    111    llii'   iiiitsiili',   till'   ii|i|>!iijitu^ 

i-N   |il,'|rril    ill    ,'l    w  iltrr  liiltli    lllllil    till'   trlll|ii'l'at  HIT'   I'lillilitiiilis   ,'111 nst.'lli! 

Till'    IllilllllllU'tl'l  N    illT'    tlu'll    I'lilMll    til    till'    nlllvillc    Jlir    mill    tllr    Imttll'S    ill! 

sliaki'ii  ill  iinirr  th.it  tlir  liriiiuL'liiliiii  Miliitimi  tliiit  is  iiiisiitiiiiilril  with 
(•.   lii;iv    t.'iki'   up  <'.   t'lTiiii   till'   ;itiiiii>|iliriT'    in   tlir   Imtlli'   iiiilM    il    lii'i'mnr^ 


ill 


'ii 


il.  J . ! 


ITf        n  l'.,ii.H.n      M I     t.i-     '1  ..! nlM.         'llil~      l,.!ill      ,.,n      :..-      H-fl     .llll.r      .1-     .-l     'llll.  ri  i;l;  . 

'! -I ut'i     'j.iyr    ,V"M    .,r    l"..r    Tin. I    im  .i-iii  niunt    nt    j.r.  N^nrf.       l'i*l     tlu     i.ltUi     |.nii'o^t-    i.iif    l.'f 

1^  r*iMn\«-T  .imT  tl;i-  pM-snrc-  r.l"  y.-is  Kiiu-KtItT  in  ll.c  Mtlu-r  liiitllf  is  iiUMMiriT  liy  llii-  Iu-ikIi'  ' 
vvlmh  It  Ills,  s  111,  ,l,.v<-  oil  ii;  III,-  ,iisl;il  tii!.,-  .>!  1  in-  iiiani.miur.  tli,-  iiunistus  iti  llic  )iii.\iM;,. 
hnili  \n-ivii  iLi'ljii-liT  1"  II-  ..nKiH.il  Kv<  1  liv  ,  ..iM|T,— .i..n  willl  llli-  l.i.i--  -iri«  ,.f  ill,'  rnljl.i  r  U:' ■ 
sll'UVIl     III     III,      I,  lilt  . . 

sfittu'.ift'il.      Tlic    rt'Miltiiis.'   sliiiiikML'c   in    tlie    vkImimc   of   tlio   atiinisplit'n 

nil  tllO  sidi'  i(f  till'  Ulikniiun  lli'llinLrluliiii  snllltinii  i'IIUm's  tlli'  I'loVC  nil 
iiioiiisi'iis  t(i  lunxc  Inwiirils  that  siili'.  llu'  ilfuiic  of  iiinvi'iiH'iit  ln'iiiLr  V<" 
poi'tidiiiil  to  till'  initial  iinsatuiatinii  nl'  tin'  lii'inn<;lol>in.  The  inanoiiii'ti'i 
tiilics  are  flicii  ai.'aiti  hrnu^Hit  iiitn  roiiiiiiiiiiii'alion  with  tin'  atinnspln'ii 
so  thai  the  iiii'iiisi'iis  of  I'lnvi'  oil  nniv  iiiom'  hark  to  its  olil  lovt'l,  ami  th' 
hottle  with  satiirali'il  lit'iiini.'liiliiii  is  ii'iiiovt'il  from  tin-  iiianoint'tcr  ami  • 
lirop  or  two  of  a  satiirali'il  soiiition  of  pntassiiiiii  t'crrii'vaiiidc  piai'ii' 
in  till'  si'parair  i-nnipaitiiH'iil  nf  tiir  liotlh'  witliniil  allowiny  it  In  ini\ 
\\illi    till-   hciiio'/lnhin.      Tin'    lioltlf    is   tlii-ii    iT-attai'liril,   tlu"   tciniii'iJiliiri 


i 

?-• 

^' 

<-  z 

a  * 

C 

ei 

CL 

1^ 
?5 

i^^ 

^ 

i 

1 

1 

1 

? 

'b 

1 

1 

c 

1 

1 

i 

cc 

c 

« 

C 

« 

•f 

4 

cS 

i 

f 

C 

4 

3 

? 

4 

t 

!** 

C 

? 

E 

?»« 

» 

0 

c 

1 

f 

<L 

« 

1 

I 

E 

? 

1 

4 

1 
1 

5 

1 

» 

« 

1 

1 

r 
1 

1 

c 

» 

N 

w 

•« 

1 

c 

* 

© 

10       30 


Penrntage  saturation 
with  oJiygrn 


too 

mm.  pnf\%t4n 


10       20      30      40       &0      60       70      80      90       tOO 

Oxygfii  prrsstiff 


|-i«      !,V        V\Vi 


mliiii;     lit    ■■\>Htii     |'!<--:it  cs    iti    H.     I't.     Jii.     4I»    aMil     lUn    urn.     mI     ttx.vmii. 
li'ir   riviit    h.ii:'l.   the  -a^iir   si-atcn   with   ttit-   (;\>K(Ii   jTrssurc  a>  ttu-  a'"*fi 
>*■  r     ly-iTt  .     .ii*'»H  i.itii'n    tiirvr     r.  iirt'.fntinji    th,-    tNiuililTiiim     !m  t^\^  i  ii    . 
ami    rt.i.ji  t  (1    lu-ii:oi;iuiiin     I  pvirplr  l.       t  Ir  Jiii    JuatpU    U.iri  r.at.) 


rol-ri  l|\rlv. 


i\    cr- 


i.\> Kt ".    o\> ln.iii"Ki'»l'in 


!        f 


i-iMM 


h    ' 


M  J 


\ 


Id -l'll(\llli\    111  ViiNI 


iillil' 


oils    ii-ii(l,jii>tci|.    tlif    iiiiilioiricti'i'    <-liiM-i|    nil'    rriiMi    th 


.111(1  the  uppanitiKt  iiitnjn  xlmk 


:i>:i 


'iiiHiilc   air, 


cii  Ml  that  the  iVii  ii-vjiniilf  mix* 


lii'iniiirliiliiii  Moliitioii.     This  iltiv.'s  off  all  the  (>,  from  tl 


ith  th 


111-  o\vhciiioirlohiii 


M>liitioii,  ami,   thfivforc,   raJMCs  the   prrssiiic   in   tl,,'   almu»|ih.'r<-  of  tl 


lioftlc  so  that   the  clove  oil 


lilt 


moves   to   the   iipiiosit)'   sill..   ,,f   tl 


Iter,   the   i\run f  ilis|ilac<'iiii'iit    liciiiir   propoitioiinl    ti 


tl 


i\yhi'mi>uloliiii 


K'   iiiaiiiiiii 

II'    illllolliit    of 


We  ha\i 


!•  now  all  the  necessary  data  for  eslimatini.'  the  rcl.itix 


c  ilmollnt^ 
ion  IIS  removed  from  the 


"''  <■<■'•< I  hciiiojfloliin  ill  the  hemiiLrloliin  solut 

tiiiiomefers,  for  it  is  plain  that  the  second  estiiiiat 

tells  IIS  how  much  oxyhemoirloliii!  miirhl   have  Ueen   furnied 


Ion,  as  discriliei]  iilmve 


moifloliin  lieeii  saturated  and  the  first 


d 


th. 


tal<en  up  liy  tl riu'inal  hemoirlol 


lUie.  how  niueh  (>,  lia.l  vet  to  li 


The  Dissociation  Curve.    Tl 

iVoiii  the  \arious  hemoKlohiii  solut 


trloliiii  solution  to  prodnc.'  s.ituiation. 
c  lle\t  step  is  to  plot   the   resiills  olitaille.l 


ions  in  th.'  f.iriii  of 


a  curve 


nowii  MS  till    ilis^ociiili 


Thi 


s    Is 


ciirrr   of  hi  ttiixihiUin.      It    is  |i!iittcd    with    th 


M'li.tive  perccntau'cs  of  reduced   and   o\y|ieiiiot;l„|,j||    j,,   ,..|,.l,   ,,,•  ,1,,.  ^„]„. 
tiuiis  iiionjr  th.-  ordiiiates.  and  the  partial  ]\r\ 


■ssiiii's  of  (»,  ill   iiiilliiiieti 


'it  III 

ihu 


eri'iiry  to  which  they  were  exposed  al.ini;  the  al 


is.'issie 


Tl 


le  curve 


s  (IrinMi   Is  exaetiv  of 


th. 


Ill'  shape  as  that    uliicli    would   he   iir 
cd  if  we  wvvi'  to  place  tlie  foiiomi.ters  in  a  row   at  distances  from  oi 


nth 


'r  eorrespondinir  to  t'le   iiaitial    pressiir..   .,f  (),    which   each 


'.lined,  and   then   to   mark   on   each    ti 


r.di 


and   oxyheriiou'lohin    t"ound   in    tl 


onoiiieter   the    relative    amounts   of 


joiiiiinr  thes.'  marks  on   the  tm 


le   dilutions   after   shaUiii!;.      A 


MH'iiil  to  the  curve  drawn  l»v  tl 


loineters    would    then    exactly 
lie  method  d.-s.-rilied  aliove.     This  wil 


"ar  from  tlie  ai-eomjiaiiyiii};  ()>;„,•,.  f,.,,,,,  Harcroffs  lnM.k  ( Kiir.  I:{7i. 
In  such  a  chart  the  space  helow  tin  .  iirve  can  lie  taken  to  represent 
■■  |>erceiita»re  of  oxyhcino-iloltin  (red  in  chart  >.  iiiid  that  aliove  it  ..f 
'iuce.l  h.'moirloliin  dilue  in  chart  i    at  th 


vliich 


ire  indicated  aloni,'  the  ahscissu'  as  1 


\aryiii'.'  p;i"i,il   pressures  of 


sphere   of   the   tonoiiiet 


crs,   and    which    must 


eiiii:  c'.i  'M.ned  in  the  at- 
lie    proportional    to   the 


ion   in   which   the  hemoi'lobin   is  di 


iiitial   pressure  of  ().   in   the  sohit 

-ilved. 

Difference  between  Curves  of  Blood  and  Hemoglobin  Solutions.  Th,- 
•irve  ohtaiiied  from  fnin  hiniDiilobiii  snhitinns  is  very  far.  however. 
'■"Ill     dearinjr     up     the     prolilein    as     to     how     the     lilo.i.l     al.s..rhs    and 

lis,-harjres     (\       ()„     tl ontrary.      it     makes     this     problem     appear 

ii  the  more  dilli.'iilt.  for.  ac.'.irdinir  to  the  curve  (Mi;  I:!")  the  heiii.,- 
••"i-m  is  Hlrea.ly  in. ire  than  half  .-omliineil  with  (>,  at  a  i)artial  pressure 
'   this  KHS  of  no  more  than   10  mm.   Ik',  which   means  that   in  the  low 

irtial  pressure  of  ().  existinj;  in  the  .apillari.'s  the  oxyhemouloliii,,  in- 


Pi  I 


I       11 


l! 


U  \l  H 


1 


I 


:!M 


I  III    ill  --I-II;  \'i  iii\ 


1^^  i 


i^l^^» 


i 


sirail  (iT  ii'ji.lilv  yi(l(|iii>;  i  its  liiml  iil'  (>.,  Wiiilld  '/li'cdilv  l-fliiill  prilr 
tii'iillv  till'  w  IhiIc  111  It.  'rill'  riii\c  ill  iitlicr  wnnis,  WDiilil  siitiNt';ict<)ril> 
•  •Npljiin  wliy  lii'tiiiiuliiliiii  •.liiiiilil  ii'iiililv  iilisoili  <>.  I'miii  the  iilvcdlm-  iiir. 
Inn    wiiiiM   I'.-ill    I'm-  sliuit   111'  cNiihiiniiiir  liow    liiis  <>.   is   r<';i<ril>-   rclciisfij 

\\  lli'll  il  is  |-ci|Uiliil  ill  till'  lissiirs.  (lliviiiiisly  tlii'li'  is  siilili'  ni'tiliriill  nili 
(litioii  incsciit  ill  tlic  ;iliii\i'  cxiiriiiiiiii!  wliii-li  imii  iml  ulilaiii  in  tin'  n;il 
mill    rll\  iliililiM'lit    III'   till'   liliiiiii. 


,00 


so 

80 
70 
60 
SO 
HO 
30 
20 
W 
0 


7^ 

^^ 

^ 

^ 

^^'^(^ 

:?= 

^ 

J 

A 

^. 

;:^ 

7^ — 

i 

/ 

// 

r 

1, 

'  i 

7/ 

V 

/ 

7 

/ 

i 

7 

L 

'i 

1 

I 

' 

^ 

/ 

to     i.0     30     HO     so    bO     70    80    *iO    100 


I  .i: 


A^.  ■,,■.■. 


I  ii-i'L.tU  -  r,  ■  .  .  i  l.ii;.     '  il'M  ..II--M    ..| 

AI-.I-.  r. „-,..„   ,  1   ,.,»„.„   ,„   n,. 

I     r  V.        V        II.  i'.  I      ..I     .,i'r,,l I     |.  ,1,      I  .  ■ 

I     IIV.      i;  l>  -I.  ^..'.I      Ihi.     ■  ,:\, 

CiHc       I  1    n,  .   I       .,1       .'O      ll,l-l        111         I,,    ..'.Til       . 

I     m      11        >'■■.      .  ,1   ■•,,U..h    .     r  V ,   M.i.,1     1,1. 


I      Till  1  .  II'  \ 

I  ul  .l-i'i    -..1,11. 


Sihcr    lli'llinijIiiliiM    t.'lki's    l|{i    (t      ill    |il'i  i|ii  il't  Ii  ill    tn    its    ilnll,    it    rilll    imt    1" 
lii'r.'HIM'    ill'    i-ll.lll'Ji's    ill     till'    •»      i-iilllliillill!i     pillt     III'    till'    llcllliiirliillill     itsri' 

tliiit  liliiiiil  Mini  |iuri'  lii'iiiiii.'jiiliiii  siiliitiiiiis  iiiiM'  ilissiniiliii'  ilissiiciiitini 
riirxcs,  liiit  r;iilii.|'  lii'i'iiiisi'  111'  ililViTi'iiccs  in  till'  I'liv  irnmnciil  in  wliii-li  tin 
lii'liKiirliiliiii  ,irls.  Tliiil  this  is  sii  IMII  lie  ii'.-nlily  slmwii  liv  plultillt;  tl;i 
(lissdriiiliiiii    i-ur\r.    iml    t'lir   a    lii'Miiiylnliin    siiliit inn,    Imt    I'nr   hhinil    itsi'i' 


RKsniJATKtN-  iii;v«.vt>  Tin;  i.i  nos 


385 


l>  ill  Fit;.  l:iH).     Tho  results  ar 


(•  very  ditVcrciit 


>(  ()..  (if  iilxnit  (iO  HUM.  Ilj;     that 


At 


ii  partial  |nrssure 


liiMf;  alveoli   (KM)  nnii.)     tho   jijood   Im 


It  IS,  a  lower  |>iessiiie  than  exist 


s  ill  the 


vJicreas  at  pressures  liel.iw  '.0  nini.  it  icadilv  loses  (»., 


leeoiiies   nearly   saturated  with  O., 


Iliere  is  I 


so  that  at  10  nii 


nearly  eoniplete  reduetioii. 
The  (piestion  is:    What  are  the  oiivironnieiital  eonditions  under  whieh 
llie  hemo!,dohin  in  the  l.loo.l  so  alters  its  eoiuhinin-:  power  for  O..  as  to 


(>nlinali>      I'lri.  nl:ii!,-    s.iliiTali.ni    i.f    Im  i.iH>;l,,l,i,i. 
Alisri-sl       T.llMnil    1,1    ijyymn    ill     Ililli.    ,i!     imniin. 
-    Hi^sMintinii    nirv,.    uf    li,in,,i;l,,l,ji,    ,|jsm.Iv,.I    i,i    , 

II,  IPisMi.i.nliiiii   .urvf    ,,f    liiiM..Kl.,l,Jn    ,li~s.,lv,  ,1    in 

III.  Ilis^..,i.iii,,i,    ,„rv,-    ,,t     li,iii.,ul,,l,iii    ,li,„Ov..|    II 
limi.ii.iliiri-    ,^;.^,^•    r.       i  I'mm     |,,„|,li     l!.,rir..lll 


Sill. 
Ml. 


;'roduee  sneh  a  dilTei  nee  in  the  disso.iation  eurve?  V,y  experinieiitinir 
Mth  heiiio(.dol.in  solutions,  three  sueh  fiii'tors  have  he.n  round  to  eoine 
■■>i'>  ])lay:  (!)  the  i)resenee  of  inor-anie  salts,  (l>)  the  hv.Irotren-i.ni  eon- 
'iilration  (CO,  tension)   of  the  solution,  and   C!)   the  temperature.     If 

'  '"lofrlohin  is  dis.solved  in  water  eontaiiiinjr  the  various  .v„/r,s  of  plasma 
■'  the  same  proportion  as  in  Moo.l  i  .ntiti.'ial  i.lasma).  (he  .lissoeiation 
'ii-ve  will  he  foun.l  to  ehani-e  so  .is  1u  resemhl,.  I  hat   of  l,lom.|    i  Kiir.    |;!()). 


i   u 


:'  1  K, 


5f     ' 


i; 


ji    •> 


f>     i  i 


I 


;t«(j 


Tin;   RKSI-IKATIOV 


Siii.-i'  tl„.  pliisiiiiis  of  .linViviil  n.ii.iii.ls  n.nli.i..  .liltV.ciit  pn.portic.ns  d 
salts,  til.,  ai'tilicial  i.lasnia  iffiiiircd  t..  scciir.-  the  irsult  is  not  always  tlir 
same.  It  ilillVis.  for  (.xaniplc,  for  tlio  .loy  aii.l  man.  I'otassinm  salt^ 
are  i)arficiilari.v  cfliciciit  in  catisint;  lifniof,'lol)in  to  alisorl)  (>..  The  in 
tluencc  of  varyiniT  liii,ln,,/i  ,i.i„n  conciiitrafiuiis  of  tlio  solution  may 
be  convonicntly  studied  l.y  a.l.iinfr  vaiyinfr  in-rcontajies  of  Co..  to  flir 
tras  mixture  in  tlie  tonometers,  when  it  will  hv  found  that  the  eurve  i.e. 
eoines  lowered  in  iiroportion  to  the  anioinit  of  CO,  i)re,sent.  This  is  shown 
in  Vlg.  140. 

The  effect  of  trmprriiturr  on  the  dissociation  eurve  is  twofold:  (1)  on 
the  rate  \\ith  which  equilil.rium  is  estal>lished  at  the  jjiven  partial  pres- 


0      10     vo     30     40     00     on     To     eo     go    ioo 

IM..    140    ■!>- ■>  ia.i....   .urv.-  „f  l,„„,anM„„,l.  „,,ose,l  ,„  0.   ;,,   ,0.  40  an.l   00  „,„.   CO..      Or.ln.a,.  , 
I  iUm      ..l,,r.,l,..„.      .\l,M,s«.i,    ,„,.„,.„    |,r.„,ir,..      .  l-r..m    .Ium,,!,    Ilar,r„ll.  J 

sure  of  0„  an.l  rl the  j.o.sition  of  the  eurve;  the  lower  the  tempera 

ture,   the  liijrher  the  i-i,rve. 

The  Rate  of  Dissociation.     Tlion-h  it  is  now  dear  that  the  thr -on 

d.tions-^namely,  .saline  content,  C,„   and   temperature-are  capable   w; 
alterni-  the  .l.ssociation  curve  of  a  pure  hemoglobin  .solution  so  as  t.. 
make  ,t  correspond  with  that  of  Mood,  this  .Iocs  not  entirelv  solve  on, 
problem,  for  w..  hav.^  yd   to  show  how  the  cooj.eration  of  these  force, 
renders   it   possible   for  the  rate  at    which   henn.trlobin   takes  up  O,   i,. 
the  luntrs  to  corrcspon,!  cNactly  with  that  at  which  it  ^ives  up  its  O 
to  the  tissues.     To  study   this  ),roblem  a  somewhat   difTerent   kind  .., 
e.xpernnent  nn.st  be  nn.lertaken.     The  hem..^dobin  solution  is  placed  in 
a  tube  and  the  ir.s  n,iNlnrc  slowly  l,ubblcd  through  ii,  samples  of  ti. 
'"'"♦'""  '"'"'^'  '■• '^••<'  ■••»  "'♦^••■V"Is  for  analysis  in  the  difTerential  blo.ni 


RISl'IKATroV    liKVOND   Till;    l.lNdS 


387 


LMs  apimiatu.s.     To  ..litaiii  tli,.  ii,tc  of  oxiiliitioii,  a  iiiixtiiro  uf  X^  or  11,, 

.111.1  (),  is  l,nl,l,l,.,l  tliroii-h  111,.  1,1 1   with  the  i)ai?ial  pressure 'of  th.' 

O,  the  same  as  that  wiiieh  ohtaiiis  in  alveolar  air— namely,  about  !)5-100 
mm.  llg;  ami  to  obtain  the  rale  of  re.liu-tion  jnire  X,  or  If,  iras  is  bub- 
Mi'd  throiijrh. 

Tiie  ratfs  of  re<liieti..n  or  of  oxi.lation  as  thus  determined  are  then 
plotted    in    eiirves   eoiistiiieted    with    the    iieireiitiiKe    saturation    of   the 


Oxidation 
n-S"  C.  no  CO, 

Hsduction 
Oxidation 

37-5°  C.  no  COji 

Reductiun 
Oxidation 

Sir,"  c. 

+  40  mm.  pressure 
of  COj 


ton 

1 

p         20         30         4 

,? 

9    ...  ?0          7(1           an          an          .«,. 

\     , 

/■ 

Rn 

y 

/ 

\ 

nn 

/ 

\ 

/ 

\ 

A 

40 

/ 

\ 

i- 

^^ 

70 

i 

"^ 

^- 

0 

L 

"                 ■ 



. 

1 

100 

80 

.r' 

— 

U- 

60 

c 

40 

\  - 

20 

\ 

-\ 

0 

'~          J 

'""— ^— = 

1 



'lriliiialf>      IVrcciil.iBi'   ^aiuraturi. 
.Miscissac. — Tinu-    in    iiiiTuiro. 
Kcducing  K.U,   hydioKin, 


l>y 


-•1.  icmpcraliire    I7.5-    C,    unit    ii..   CO, 
'',  teniperaliirr   .17  5'    0.   with    ih.   lO." 


€ 


<'.  teiiiperatiirr   .17,5 
'•■^(•pli  Harcroft,) 


I-,.    I,„l    Ihv    ..,   ;„„1    II    ,„n,ain.-,l    w   nM„.       llg  |,r,.,«„.e   nf   CO,,,      ,,:,,.„ 


pr 

-1 

"   ■: 

'  f 

;t: 

-| 

4 

.   1 

388 


Tin:  Rrsi'iRATioN 


^•111 


If 


licmodldliin  (iM  t!ic  (Hiliniilcs  and  tlic  tiiiic  in  niiriutcs  iilonj,'  tlio  absciss;!' 
(V\g.  141 ».  KvfM  if  we  use  i)liMiil  in  tiiis  rxin-riincnt  and  tlicivfori'  iiiaki- 
•MTlain  tliat  the  licninirlnliin  is  aclin-r  in  tiic  jjicsfncp  of  tlii'  iiroppr  pro 
I>ortioM  o;'  salts,  wo  shall  (ind,  as  Kiu'.  A  sliows,  that  at  ntoiii  tcinpcratuif 
till'  lalc  of  oxidation  is  very  iiiucli  creator  than  the  rate  of  rodupfion 
If  now  UP  ri'iipat  tlic  ohsprvation  at  a  tpnippratuip  of  ;{7"  C,  the  two 
piirvps  ponip  more  iipnily  to  poiifsijond.  Imt  still  tlip  ratp  of  rodiiPtion  is 
slowci-  than  that  of  oxidation.  If  in  a  third  pxpprinipiit,  hosidos  hnvini.' 
l)rop('r  tPMiiipiatiirp  and  plipiiiical  conditions,  wp  prodiicp  the  oxidation 
and  reduction  in  the  iircs"ncp  of  a  jiartial  itrcssurp  of  CO.  of  40  mm., 
Mhich  coii('s|i()nils  to  that  of  the  attciial  hlood,  wc  shall  find  that  oxiila- 
tioii  hcconics  a  little  slower.  \v  heieas  rc(lnctioii  is  fnrtlicr  (luickpiipil 
Fmhcd  the  two  cmi\('s.  as  seen  in  T  in  the  liuuip,  poiiw  i)ractipally  to 
correspoiid.   imiii-jitinii-  that    the  environmental   conditions  undor  which 

hcnio'.'lohii iidiincs  and  <ri\es  olT  O,  in  the  Mood  are  exactly  ad.jnsted 

One  word  inon-  with  rc'.'ard  to  tiie  inlhlencc  of  ('„.  Its  cfTcPt  in  flat 
tPidnj;  out  the  cur\c.  especially  .-it  the  lower  jiartial  prcssiirPs  of  (>  , 
indicates  that  when  a  hiudi  C,,  is  pn'sent.  the  Mood  will  very  readily  part 
with  its  <>,  supply.  \ow,  the  most  si'.Miidcant  application  of  this  fad 
is  that  liiu'Ii  concentrali(Uis  oi'  II  inn  will  occur  .just  exactly  whprp  if 
will  hp  of  henclit  nauicly,  in  the  ea))illarips  (liocansp  of  thp  CO,  and 
lactic  acid  prndui-cd  liy  the  tissues").  Sonic  doiiht  has,  liowpvpr,  rpppntl.\ 
heen  lliinw  n  on  the  importance  of  this  factor. 

Since,  as  wo  ha\e  scoi.  henioLrloliin  alisorhs  (>,  accordinjr  to  phpmi<'al 
laws,  it  will  naturally  he  askcil  not  only  why  the  dissociation  onrvp  flat 
tpns  out   while  yet   iiiaintainin'r  the  shapp  of  a  risrht-ansrlpd  liyperliola. 
as  hy  tlip  action  of  ;icids  or  an  increase  in  tpnii)erat>irp,  hut  also  why  il 
should  ohansrc  its  shapp  when  salts  are  also  iircseiit.     The  explanation 
ofTercd   hy   IJarcroft   jiiid  his  jiupils  is  that   the  chanijps  dppond  on  the 
fact  that   hemou'lohiu  heiiiu'  a  colloid.il  suhstaneo,  its  molpoiilps  lindpifrn 
procossps  of  au'irrcuatioii   under  the  conditions   rcferrpd   to  aliovp,  aiul 
therefore  cause  the   reaction   to  hccome  of  a   difTcrpiit    typo  from   thai 
rciiresented  hy  the  equation  llh().;=illli     (»,.     As  has  lipon  pointed  out 
hy  Hayliss.  althouirh  such   an  explanation  niit;lit   sufiicp  to  pxi)lain   the 
flatteiiinir  out  of  the  ciirvp.  if   f.'iils  to  exiilaiii  the  phanjip  in  its  shape; 
for.  accordiiisr  to  the  laws  of  m.iss  action,  such  a  chanjrp  could  opciii 
only  if  niolcciilcs  of  a  difl'eicnt  type  came  to  takp  part   in  the  rpaptioii 
Dissociation  Constant.     N'ofw  itlistandiu!;  tlipso  priticisms,  it  is  of  pon 
sidprahle   lu-actical    imiiorfancp   to   know   that    an   pfpiation   pxists  from 
which  the  Piitiro  dissociation   ciicvp  can  he  plotted  hy  m.-ikins;  onlv  nu: 
determination  of  tho  rplativp  amounts  of  oxy-  and  reduopd  lipmoKlohin 
at  a  particular  tension  or  partial  pressure  of  oxy-rcn.    This  oipiation  is  a- 


Ki:si'IHATH)\    DKYOM)   Till;    I.l  \(iS 


389 


r.>ii( 


KM) 


1  I  K; 


fir  I)  tMiuals  the  lu'ivi'iita'.'f  satuiatioii  of 


li.'iriipj;l..liiii  with  ()_„  .  till'  <>,  fciisioii,  and  l\  aii<|  ,i  air  coiistjints.  A" 
lii'injr  tlip  ciMiililn-iiini  (•(instant  and  ii  the  avi'ni;;('  iiuihIht  of  nioKrulcs 
(if  li('inoj,'lol)in  supposed  to  exist   in  cacli  a jrtr rebate. 

When  tills  ecjuatioii  is  applied  to  hiiiuan  lih.od,  tlie  value  of  ii  remains 
iiiiehan<;ed  and  is  niveii  as  'J.'),  so  that  liy  tiaiis|)c,sitioii  we  are  eiialiled 

to  find  the  value  of  K  as  follows:  K      --;-,„i^.    -     .     II'  we  lind  the  value 

X'  (  Kill      y  ) 

of  K  liy  iiieasiirinjr  the  relative  saturation  of  the  iilood  with  O,  ntoiii  pres- 

Mire  of  tiiis  <:as.  then  l>y  ehanniii','  the  value  of  x  to  correspond  to  other 

<>_.  pressures,  we  ean  lind  all  positions  of  the  eurve  for  a  ^'iveii  saiiiiile  of 

Iilood. 

An  important  praetieal  applieation  of  this  metliod  is  found  in  tin 
lUhii.ihialion  of  Ihr  ('„  i,f  hl„(„l.  for,  as  we  ha\e  si^eii.  the  dissociation 
curve  iieeoiiies  lowered  in  proportion  to  the  concent ral ion  of  liydioijcn 
ioiis.  The  acidify  of  a  sample  of  Mood  .an  theii'fore  lie  found  liy  com 
parison  of  its  dissociation  eurve.  as  plotted  from  the  values  found  for 
A',  with  fliaf  of  normal  Iilood  to  which  known  i|iiaiitities  of  acid  have 
licen  added.  Wlien  the  curves  correspond,  the  liloods  iiinsf  contain  the 
same  amounts  of  acid,  other  thinu's  lieiiwj  ei|ual.  In  Inicf,  then.  Ihr  rr- 
'irliiiii  of  Ihr  hhiml  is  i>roi).)rtii)iiiil  lit  Ihr  rnliir  of  A'.  When  this  is  low. 
it  inilieates  that  the  Mood  is  takiiis;  up  an  alniornially  low  pereenfaire 
i'(  its  possible  load  of  0_,  at  a  iriveii  pressure  of  ().,  and  that  the  acidity 
IS  <;reafer  than  normal:  when  A'  is  liiirli,  for  the  same  reason  the  acidify 
iMUsf  be  low. 

In  defermininu'  A'  for  the  hjood  as  it  exists  in  the  l.ody.  it  is  i essary 

that   it  .should  he  siih.jected  to  the  same  tension  of  .         as  obtains  in  the 

■'' 1  vessels.     A"  will  then  be  proportional  to  the  C,,     ,  the  living  blood. 

This  condition  would  be  impossible  to  fiillil  in  drawn  samples  were  it 
not  for  the  fact  that  we  can  place  in  the  tonometer  an  atmosphere  eoii- 
iMiniiii;  the  same  partial  |)ressure  of  Cn.  as  is  found  in  the  alveolar  air. 
Since  this  valu"  varies  in  dilTereiit  individuals,  it  must  be  separately 
•  iscertaineil  in  each  case  i  see  p.me  ■.'Ai\.  .\s  d.terniiiied  »itii  these 
iiiodilicafioiis.  A'  has  been  found  to  vary  in  healthy  men  between 
IIIH»0212  and  0.00(t:t(i:!  (ten  individuals^.  When  acid  substances  appear 
111  the  IiIoim],  as  in  acidosis,  A'  bei'omes  extremely  low:  tlnis,  in  one  ease 
Mifferintr  from  acidosis  with  dyspnea,  it  was  found  a   few  hours  before 

death   to  be  only   from   O.OddOS'J   to  0.0(1011.     .Similarly   A'   Ik iiies   low 

111  the  acidosis  associated  with  iiiounlain  sickness,  and  it  is  .said  to  be 
raised  after  taking  food  that  is  rich  in  alkali.' 


'Whrn  A-   is  fmin.l   I..  I.,-   n..rn,.,l.   lli,-  l,h...,i   i,  .,,i,l   l..  I.,.   „„•„■,  (,V.    uli.r,-    K   U   l„„     it   i«   v,i,|   i,, 
11  mvont-rd,  ,■  ami  wlu  ii  K  is  Inijh  an. I  llu-  ,i<  i.hly  is  tlurvforc  small,  ll  is  sai.l  t,.  I,..  fl,„ii,\lU. 


•  isl  > ' 


jl  ^ 


I  pi 


^ 


4i 


VUAVTEli  XLV 
Ki;si>II{ATI(i\  IIKYOM)  TmO  l.lNdS     C.ntM 

THE  MEANS  BY  WHICH  THE  BLOOD  CARRIES  THE  OASES 

111  till'  luivKoiiiir  ;i.v„unt  .,t'  \\w  iili.vsidlouy  <.f  tlio  IjI.mp.I  fjases,  (.iiiplia- 
sis  is  placed  dm  tlif  tiusiMii  uihI.t  wliicli  Ww  tras.s  exist  ratiior  fhiiii  on 
the  tutal  iuiicMiiit  (.1'  each  -as  |.ivmmiI  in  tlie  1,I,mm|.  Tliis  has  been  done 
i'eealise  the  exehaiitre  of  -ases  lietweeii  alveolar  air  and  I.lood  and  1m - 
tueeii  hi.iod  and  tisNues  i)roeeeds  aceordin^r  to  llie  laws  of  i;as  dilTiision. 
uliieh  aie  oi'  course  dependent  upon  dilVerences  in  ^iis  pressure  or 
tension. 

Sonietiiinu'  must  now  h.'  said  reL'ardinjr  the  nininnil  of  Ihr  tinani.  This 
may  he  measured  either  hy  i)h.\  sical  itv  h\  chemical  metliods.  In  tlic 
former,  a  measured  <piantity  of  lihiod  is  received  into  an  evacuated  >;lass 
vessel,   wliicli  is  then  attaelie.l   to  a   mercury   pump,  liy   which  tlie  irases 

are  suck.d  out  of  the  hi 1  ;ind  transt'erred.  hy  suitahle  manii)ulatious 

of  stoix'oeks.  to  a  irradiuited  tuhe.  in  whicii  tlicy  are  then  analyzed  li.v 
ehenncal  means.  The  principle  of  the  chemii-al  method  has  already  Iieen 
descrihed  in  eoiniection  with  the  measui'enient  of  oxyi^eu  in  hemonlohiri 
solutions  (see  pajre  :!S2i.  A  iiu'asured  ipiantitv  of  hloo.l,  kept  ivi','  from 
contact  uith  the  air.  is  transferred  under  some  weak  ammonia  solution 
to  one  (d'  the  hlood  jras  hottles  of  the  Mood -yr.-is  ilitTcrent ial  maiionu'ter. 
and   a    U'W  drops  of  a   saturated  solution   of  potassium    ferrieyanide  is 

Iliac. •({  in  the  i)ocket  of  the  liottle.     After  the  hi 1  has  heen  hiked  anil 

temperature  conditions  ;id,juste.I,  the  ferric.v  anide  is  ndxod  with  the 
hlood  solution,  thus  catisini:  the  O..  to  he  ipuintitatively  displaced.  From 
the  incre.'ised  prcssurr'  pro.liiced  in  the  manometer  the  amoinit  (d"(>.,  can 
re.adily  he  couiputcd.  To  determine  thi'  Co,  of  the  hlood.  the  bottle  is 
now  removed  from  the  manoirieicr  and  a  few  drops  of  a  saturated  solu- 
tion <.f  tart.irie  a<'id  iilaced  in  the  pock.^t.  When  this  is  mixed  witti  the 
deoxyt'emited  hlood  mixture,  after  the  usual  ad.ju.stiiient  for  tempera- 
ture, the  pressure  eau.se<l  hy  the  .•\ol\ed  CO,  is  recorded  and  the  amount 
present   calculated. 

The  results  of  the  analysis  are  ex|oessed  as  the  nnmher  of  euhie  eeiifi 
nu'ters  of  ^Ms  present  in  KM)  ,•.,•.  ,,f  hlood  the  volume  i)ercentaf;c.  as  it 
is  called.     The   followinir  ••ire   approximate   percenta;;e   values: 


1*1 


RKSI'IKATION    IlKYOM*   TllK    I.IMiS 


:jyi 


Venous  Mom] 
Artorinl    Mood 


12 
20 


CAKBON'   UIOXIUC 

48 
40 


TOTAL  GAS 

60 

60 


The  estimation  of  tlie  ainounts  of  the  sjascs,  although  of  little  value 
in  ponnoction  with  the  ])hysiolo(jy  of  gas  pxchanjio,  is  very  important  in 
sii|)i)Iyin(r  information  rotranlinir  the  rcspii'atory  activities  of  the  various 
ortrans  and  tissues.  Just  as  we  determine  the  total  respiratoiy  exehange 
of  an  animal  liy  measuring  the  dilTereniM's  in  (^.  and  ("O,  in  inspired  and 
expired  air.  so  may  we  dotermine  the  <lei;ree  of  tissue  respiratory  ex- 
••liaiige  by  analysis  of  the  gases  in  hlood  I'emoved  from  the  artery  and 
vein  of  the  tissue.  It  should  be  elearly  understood,  liowever,  that  it  is 
not  the  percentage  but  the  total  amount  of  the  gases  that  must  be  eon- 
sidei'ed.  and  tiiat  it  is  therefore  necessary  to  know  the  volumes  of  blood- 
How  as  well  as  the  ijcrcentace  of  the  gases.  Something  will  be  said  later 
of  the  results  (if  such  in  vest  i'.'at  ions  (see  jiage  :!0'>). 

At  present  we  are  eoneerned  with  the  manner  in  which  irases  are 
carried  in  the  blood.  The  0._,,  as  we  have  seen,  is  carried  by  the  henio- 
L'lobin,  some  being  also  in  a  state  of  simi)le  solution  in  the  i)lasma.  The 
>'0..,  wliich  it  will  be  noted  is  present  even  in  arterial  blood  in  eon- 
side  .tbly  greater  amount  than  the  ().,  is  partly  combined  with  alkali  to 
form  bicarbonates.  The  alkali  available  for  this  ])uriiose  varies  from 
time  to  time  acciu'diiig  to  the  amount  of  other  acid  substances  present. 
Since  these  are  stronger  acids  tluin  carbonic,  any  increase  in  their 
iiniount  (acidosis)  causes  displaeeuKMit  of  some  of  the  CO.,  tluis  bring- 
ing about,  as  we  have  seen,  a  relative  increase  in  free  CO.  in  the  blood 
;ind  therefore  raising  the  {',|. 

Wliat  particularly  interests  us  here  is  (lir  iifiniifi  hi/  wliicli  the  com- 
hiiird  CO..  Is  rnrrifil  in  tlir  hloail.  If  lilood  is  exposed  to  a  full  atmos- 
jihere  of  ("(V,.  it  will  take  up  as  much  as  LW  per  cent  of  the  gas-^that 
is,  between  two  anil  three  times  the  anioinit  or<linari!y  i)rcsent  in  it. 
It  has  therefore  a  great  reserve  <'apacity  for  CO,..  A  srieater  i)ropor- 
tion  of  the  CO,,  is  carried  it;  tlie  i)hisma  than  in  the  corpuscles;  but  if 
plasma  Cor  seriuii )  is  exposed  in  a  vacuiim.  all  of  the  C<1_,  |)resent  in  it 
will  not  be  evolved.  When  !)lood  itself  is  similarly  expo.sed,  on  the 
other  hand,  all  the  CO.  is  iriven  off.  To  liberate  all  of  the  CO,  from 
plasma  in  raiuo,  some  acid  must  be  added,  fnmi  which  it  has  been  in- 
ferred tliat  blood  eori)uscles  act  like  weak  ai-ids.  It  is  eommoidy  stated 
that  hemoglobin  or  some  eonstitui-nt  of  blood  is  capable  of  freeing  CO,. 
from  solution.s  of  sodium  carbonate.  l)ut  the  recent  work  of  Buckmaster'- 
>lio\\s  that  this  is  not  the  case.  The  decomposing  power  of  blooil  is 
caused  by  the  devi-lopmcnt  of  acidity  in  the  shed  blood  and  any  similar 
[lower    that    the    corpuscles   may    exhibit    is    due    to    a   discharge    from 


Illi 


!j  J 


hill 


i  r 


392 


tin;  kksi-ikation 


tlu'iii  into  tilt'  pliisiiiii  i<(  iicid  liidiclcs.  At  Icnst  it  has  In'cii  t'oiiiiil  tlmi 
thf  iilkiiliiiily  i<(  the  pljisiiiii  iiKMciiscs  wlii-ii  CO.  is  Ituliltlfd  tliroii;;), 
Iil(i.»I,  this  iiicn-iiNC  in  alkalinity  licinu'  inlcrprflc.l  as  llic  n-sult  of  lli. 

'"''-''■•'"i •"  '•••i<l   fa<lii-lcs  into  the  coipiisclcs.     This  would   lead   us  t.. 

<'X|M'i-t  that  iiiidi'i-  thf  opposite  conditions  (i.e.,  in  rnnio)  acids  woull 
leave  the  corpuscles. 

Troteins  are  amphoteric  suitstanecs  that  is.  they  condiiiu-  with  acicN 
or  alkalies  which  would  lead  us  to  expect  that  they  woidd  he  <-apahl. 
of"  alisorl.iu-r  some  CO,.     That   tliis   is  the  cas<-,   particularly    for   lician 


~   70 

J  ** 

c  60 

4  50 


o 


45 


,  40 


y^ 

>^ 

».^          ^^ 

^^          ^^ 

^2  ^^     " 

/5^ 

At^      ' 

^-Z^    ^^ 

/ 

J    -' 

-  V— 1 

^^ 

/  / 

/ 

7      r 

, '                '" 

/     , 

J. 

?      ' 

T 

.{_ 

30         40  10         60  70  80         90 

''^     "'       *"•>'■   "I    <" : I.      l-..r   ,|..  ,,|.t,..„.   -.,    ,>M.      .l-r.„„    ll,„„.a.,-,„,    II.,,,.. 

I..S  .!■.  I  n,ii.i.,i.,  ,1 

«■'"'""•  'I'l^  ' 11  shown  hy  comparin-  the  CO.coud.ininfr  (M.w.'rs  of  water 

ami  a  solution  of  pure  hcimnrloliin. 

Attempts  have  lieen   made   to  determiru-  the   relative  amounts  of  CO, 
carrie.l  |,y  these  various  ajrencies  in  the  l.lood.     The  followinj;  is  an  e.\ 
ample  of  such  a  tahlc; 

III  sini|,Ii-  >.,lijih,ri   III  |.|;iMii:i  :iM,|  ciipusili  .s  l.ii  ,.,,._ 

\  r...      1  1         .        I  a   in  im  inisclrM  (is  I 

.\>    ■i.i.liiiin    l,h':iil„,ii:i  |.  '  iss     11 

I  ,1  III   |,l;isiiia  l:;.(l  I     '^^ 

III    ' l.ill.ltiiin    with    ll(.|lh,;;l(,l,i|i  7.-,    I 

I Miil.iiiiiliiih    with    |,i,,li.ins  ,,)■   phmnni  ]  1  .s    (     '"•' 


(Locwy.) 


40.0 


RI->i|-IKATI()N    IlKVOND   THK   I.I  MIS 


393 


The  power  of  lilood  to  nlisoil)  (*()j  at  various  tensions  of  this  kiis,  ns 
ilftcriMJiU'd  ill  the  siiiiic  \v;i.v  as  for  ( >.  (wr  ]m\n'  :{M()),  ha.s  shown  that 
siitiiration  of  the  )i(>nioi;hil>in  with  (),.  distinctly  diminishes  the  CO,.- 
canvintr  power  of  the  lihioil.    This  is  shown  in  the  neeonipaiiyiiic  eiirvos 

The  various  tensions  of  ('(»_,  are  (fiveu  ahniR  the  ahseissiu  and  the 
voiuiiie  i)er  eeiits  of  CO.  tal<en  np  liy  the  blood  on  the  ordinates.  The 
ujiper  eurve  is  drawn  from  results  obtained  wlien  tlie  lilood  was  shaken 
with  CO,  in  fhe  i)rese!iee  of  liydrojren,  and  the  lower,  when  in  tlie 
]iivsenee  of  air.  (The  dotted  eurve  may  be  disre«arded.)  The  line  Alt 
iliawn  between  Mie  two  enrves  represents  the  absorption  of  CO,,  by  the 
lilood  within  the  body.  At  a  tension  of  40  iiiiii.  CO.  -that  present  in 
.ilveolar  air  (see  pa<rc  :(.">»;)  .1  st^inds  in  arterial  blood  at  about  .">2  vols, 
fur  eeiit:  and  at  a  pressure  of  (>2  nun.  possibly  present  in  the  tissues  - 
/.'  stands  in  venous  blood  at  about  (i7  vols,  per  eeiit.  The  CO, containing' 
power  would  be  7  per  eeiit  lower  (i.  e.,  fiO  vols.  i)er  eeiitl  in  blood  saturated 
with  O.  at  the  latter  pressure.  The  oxy};<'nation  of  blood  in  the  liiiit;s. 
therefore,  helps  to  drive  out  the  CO.:  and  eonversely.  its  deoxyjienation 
ill  the  tissues  enhanees  its  power  of  absorbing;  this  fins. 

Havintr  shown  how  the  blood  transports  its  eharire  of  O,  from  the 
liinjis  to  the  tissues,  we  may  now  iiroeeed  to  study  the  eall  for  O.,  by 
the  tissues,  and  in  this  eoniieetion  we  have  to  eonsider  (1^  the  amount 
"f  O.,  which  they  re(|uire  under  varyinjr  conditions  of  rest  and  activity, 
and  (2)  tlie  meeliaiiisms  by  which  their  varvin<;  demands  are  met. 


i 


r 


1 


THE  OXYGEN  REQUIREMENT  OF  THE  TISSUES 

Tn  orilcr  to  ascertain  the  avcrat'e  O.  rei|uirement  of  the  difTereiit  tis- 
Mies  of  the  body,  it  is  ni-ces.sary  to  ailopt  as  a  standard  of  measurement 
tlic  aiiiouiil  of  O.  in  c.c.  absorbed  per  stiiim  of  tissue  ficr  ininiite.  To  ob- 
t.iiii  it  we  must  know:  (11  the  wei<rlit  of  the  particular  or};an  lu-  tissue 
under  invest ijrat ion;  (2)  the  bloodflow  throujih  the  vessels  of  the  organ 
ill  c.e.  per  minute:  and  (H)  the  dilTcrcnt  perceiitasres  of  0„  in  the  arterial 
.iiid  venous  bloo<l  of  the  ti.ssue.  It  would  be  beyond  the  scope  of  this 
liook  to  review  in  any  detail  the  many  ex|)criniciital  investijrations  which 
liavc  been  undertaken  in  this  connection.  A  few  of  the  most  recent 
:iinl  important  results  arc  -riven  in  the  accompanyin<j  table  from  llalli- 
liiirtoii  's  /'/ii/.v /((/<*(/(/ : 

111  the  order  of  their  oxy>rcn  re<piirements,  or  tlir  roifficiriil  of  o.r'nhi- 
liiiii,  as  it  is  called,  the  tissues  may  be  divided  into  four  jiroiips;  friandular, 
;iiiiM-ular,  eoniu'ctivc,  and  norvnu.s.  The  nervous  lis.sues  slioiild  jiossilily 
4and  above  the  eonneetive,  but  very  little  is  known  refjardiuK  their 
■  i.xyRen  consumption,  althiuifrh  it  appears  that  this  is  (piite  low  (Hill  and 


i,. 


1 

11 

1 

'M 

' 

"'; 

!■' 

i  n 


t,  ' 


i  \ii 


i'1 

I:! 


;*'' 

^^ 

.  ,  I 

• 

i 

:  ' 

= 

lit 

1; 

.'I'M 


THK   KKSI'IKATION' 


oRdAN  coMirnnx  Of  rfst 


oxvnrv  fsci) 

I'KR  MINI  rr, 

ri:n  (ifiAM 


COMIITIOX   OK    AlIiVlTY 


oxvors 

I'SKD    P.R 

MINTTK 
VVH    fiRAM 

Of   niicAN 


Viilmitary  N'irvcs  cut.    Tunc 

liili;«rli'      ■  uliscllt 


I'liHtripnl  K.-ttini; 

lullsfic 


lli'urt 


Very  slow  ntnl 
firlili'  (•(iiitriictions 


Siil.innxillarv       NVrvi-s  cut 
alriml 


i'nni'icns 

KiillH'V 
[lltCstilHW 

l.ivcr 


N'nt    -.('iTotin;,' 

Si-iuity  socrotiiin 

\i>l  ;ili>iuil,iii(f 
III  ^ll^tin^  iiiiiiiial 


SiipnuiMinl  Xnrinnl 

::liiiii| 


n.lio:i  cr.       Tunc  |.xi-.liiiir  in   ri-st 
•  inill.'  CMntMii'lioii 
.\cli\r  I'iMiti.'ii-tioii 

IMWJ  CO.       f'l.iilriutiiiB 

0.007  cr.        N'"rm:il  idiitiai'tionn 
ViTv  iictivo 

O.o:!  cc  Chnrila  stiniiilatinns 


0.(1.'!  I'.c  Sccrotiiiii  iiftiT  iiiji'i 

lion  of  Hciictin 

O.li;!  cc  Al'lcr  in.j.'ction  of 

llillM'tic 

O.d:;  cc  .M.siiiliin;;  |>c|itMni' 

0.01     to  111    fill    llllilllill!) 

O.OJ  cc 

0.043  cc  


O.OOIl  |.  ,• 
o.ii-O  e.t'. 
O.IISO  IM-. 

O.no;  r.r. 


O.O.")  cc 
O.OH  cf. 

o.in  ,..c 


O.IO   |..p 
11.10   cc 

o.n.i  p.c 
o.o::   t.p 

0.0,-|  cc 


XaliaiTol.  It  is  of  coiirso  iioccssiu-y  in  iiiakiiij,'  tlicsc  (•(imparisons  to 
sccuir  tlip  coefticiciit  of  oxidation  l)otli  when  the  tissue  is  at  rest  an. I 
when  it  is  thrown  into  varyini;  ileirrees  of  activity.  Siiecial  attention 
lias  lieen  devoted  to  the  re(iuin'ments  of  sUeh'tal  niiisele,  heart  nnisel. 
and  tlie  salivary  glands. 

Skeletal  Muscle.     In  ol)servations  on  skeletal  muscle,  Verzi'ir  (ef.  27 
i.solated  the  frastidenemius  niusele  of  the  eat,  and  without  distufliin-j  it- 
Mood  supply  eolleeted  .sanii)les  of  Mood  by  intfodueini;  a  1   e.e.  jjipetti 
into  a  lifaneh  of  the  saphenous  vein.     Activity  was  produced  liy  throw- 
in-r  the  niusele  into  tetainis  liy  the  application  of  .in  electrical  stimuliiN 
to  the  sciatic  nerve.     Duritifr  its  contraction  the  muscle  lifted  a  weiRht. 
so  that  it  did  almut  70  jrrani-eentiineters  of  work  at  the  tie!:inniii<r  oi 
each  period  of  tetanus.     The  velocity  of  l.loodtlow  was  determined  li.\ 
the  rate  at  which  the  Mood  (lowed  alonjr  .he  pipette,  and  the  O,  consiiiiii. 
tion.  by  the  difference  in  percentajre  of  O,  in  the  venous  and  the  arteri.ii 
Mood.     These  measurements  were  made:  (Ji  before  contraction,  (2)  diir 
inp  contraction,  and  C!)  after  contraction.    It  was  found  that  during  tin 
tetanus  the  0.  consumption  in  some  eases  was  greater  than  durinf;  rest, 
while  in  oth.'fs  it  was  actually  les.«,  but  in  every  inst/uice  .-i  ^'r.-jit  incr<'n>-^ 
in  O,,  consumption  followed  the  tetaiiii.s     that  is,  the  call  f.n-  O..  continue- 
for  .some  time  after  the  actual  work  has  been  performed.     This  resu'i 


RIXI'IRATION    IlKYONI)   Till;    I.INdS 


:i!tr> 


shows  tlint   flic  colli riK-t ion 


the  oxidation  ocnirit  after  the  contraction 


is  not  (lc|n-ii(lciit  upon  oxidntion,  lnit  lliiit 


i»  over.    The  nioclinnisiii  involved 


ill  niuspuliir 
iliich 


contraction  ciiii   not   tticicfore  he  anuloKous  with  that   by 
li 


which  rnprjfv  is  lilicnitcd  in  a  steam  cnu'inc  iiy  the  oxidation  of  the  coal. 
The  mechanism  must  rather  he  like  tlint  tit'  a  sprini.'.  wjiicli  liecomcs  iin- 
uoiind  diiriin;  the  miiseiilar  conlra.'tion  and  re<inires  O^  for  its  rewindiiii.'. 
Tnterestiiiu  results  corroliorative  of  these  conclusions  have  l.eeii  se- 
cured by  observations  on  the  Imtl  iinnhirliiin  of  isolated  muscles.  It 
was.  found  that  licat  production  occurred  after  a  sinifle  shock  to  the 
iriuscles.  not  only  durinir  the  contraction,  but   for  a  eoiisid<'rablc  period 

after  it.  jirovidcd  O,  was  present.     In  the  absenc  of  (I.   this  r .very 

uas  either  greatly  delayed  or  entirely  abolished.  Such  results  favor 
the  view  that  (>,  is  used  larcely  in  the  processes  whereby  the  muscles, 
"like  an  eiiyine  charirinu'  an  accuiiiulalor,  synthesi/.e  substances  eon- 
tainin«  a  considerable  aniount  of  potential  eiierfix.  which  ai:ain.  like  the 
accumulator,  it  discharires  when  appropriate  stimuli  are  applied"-  (I,. 
\  Hill.  cf.  27).  One  immediately  thinks  of  lactic  a.-id  in  connection 
uith  these  inteiTstiiij;  results,  for.  as  has  already  been  stated,  Hopkins 
iiid  I-"letcher-''  have  shown  that  this  aci.l  is  priMluced  in  III.'  abscn.'e  of 
(»,  in  excised  fro«  miiscles,  but  when  0_.  is  j. resent,  it  is  either  not  pro- 
duced or,  if  so,  quickly  disappears. 

Heart  Muscle.  Another  muscle  that  has  been  thorouirlily  iuvestiirated 
ill  this  connection  is  that  of  the  heart.  The  aaseoiis  exchanire  has  been 
.studied  both  on  isolated  hc.irt  preparations  ami  by  exainiiiiii<j  the  ex- 
chaiiire  in  the  lunj.'s  of  a  combined  luiitr  and  heart  i>reparation.  Tlie 
most  important  investigations  by  the  first  of  these  methods  are  those  of 
liolidc  (cf.  '!'.  who  arrived  at  the  very  important  conclusion  that  the 
(1.  taken  in  by  the  heart  muscle  varies  directly  with  the  maximal  ten- 
sion set  up  in  the  heart  by  the  coiitiacijon.  This  tension  was  measured 
liy  placing'  a  rubber  ba;;  in  the  ventricle  .nid  disteiidiii','  it  with  water  at 
a  known  pressure.  V>\  jiltcriiif;  the  initial  iircssurc  and  by  observing;  tlie 
pulse  rale,  it  was  found  that  the  ( ),  used  by  the  heart  depends  on  the 
pnidiict  of  the  imlse  fieipiency  and  the  niaximal  increase  in  i)ressiire 
luodiieed  by  each  cardiac  contraction;  oi'.   in  the   lorm  of  an  euuatioir 

Q 

—  =  a  con.stant  (|uantity:  where  Q  is  the  owu'en  used.  T  the  maximal 

NT 

increase  of  jircssure  at  each  beat,  and  \  the  frcipiency  of  the  ]Hilse. 

It  should  be  pointed  out,  however,  that  constancy  in  the  p  •oduet  of 
itie  nbiivo  fi|Uatinii  lioes  not  hold  under  abnormal  conditions  of  the  lieart- 
bcat  For  example,  when  the  iiressure  in  the  lieart  is  very  hiirh,  the 
amount  of  O.  rcipiircd  bcfrins  to  sjo  up  out  of  proportion,  indicatiiij;  that 


Ft 


=  •1 


II 

-•jfi 


iff 
ft. 

i 


:i!Hi 


TIIH    KKSI'IKATHIN 


\\ 


#i/' 


r  ■ 


:f 


!■  1 

■ 

':; 

^ 

III 

%m 

,V.k 

tlir  lii'iiil  is  lii'i-iiriiiiiu'  ii\i't'tii\>'il  lliiit  it  i>i  I'isiiiif  its  rfllfieiicy.  Tl' 
MiitD'  rrNiilt  iii-riii'H  wlii'ii  till*  Ix'tii't  ix  il\  iiiK.  aiitl  wlioii  (IfprcHHiiii;  iIiiil 
iiic  used,  Niii'li  iiH  rliliii'iil  liyilratc.  |)<itiissiiiiii  cviiiiiilt*,  vt'nitriiu',  it' 
Siuiic  dtlici'  ilruifs,  liiuM'viM-,  siicli  IIS  I'liiiK-iiliriiif,  ilo  not  cause  iilfci. 
liiiii  ill  tlir  iiitiii,  iiin'  lilies  Njimis  stiiiiiiliiliiiii.  Of  ciMii'sc  wlii'ii  tlir  vau'ii 
is  sliiiiiihiti'il,  till-  (^  i-iiiisiiii<|itii>ii  ill  ii  v'ivrii  piTiiiil  ilci-irnsi's  lii'i-aii-" 
till'  lii'iirtlii'ats  arr  slnwcil  ;  liiit  tlir  iilisiirptiiill  nl'  <^  is  lint  iliiMi'llscil  rv\u 
li\i'l\    III  tli-'  slow  ill".'  Ill'  till-   liriilt. 

Olandl.  Must  work  has  iiallllallx  lici'ii  iliMic  mi  llii'  iinst  iii>i>(>sNilil' 
U'laiiil  till'  siiliMiaxillarv.  Ity  stiiiiiilatiiiu  tin-  siTi'i'tni'v  iiitvc  "f  tin 
ulaiiil  itlii>  cliiii'ija  t\iii|iaiiii  in  Ilii'  ijiii.'.  it  has  liii'ii  I'iiiiih!  that,  whi'ii;! 
till*  iiini'f  aliiiiiilaiit  sri'i'ftioii  liiNts  iinlv  so  liiiid  as  tlu*  stiiiiiiliiH  is  i\\< 
plicil  to  till'  iii'rvf,  till'  <•,  i-oiisiiiMpliiiii  is  iniTcasi'il  to  scvcnil  fiiiii's  tlin' 
III"  ri'st.  ami  i-i'iiiaiiis  ini-i'i'aM>il  for  a  ri.nsiiliTalili'  prriml  al'liT  tin'  stiiiiiiln- 
has  lici'ii  irliiiiMil.  Ai-rii|iipaii.\  iiii;  llir  iiii'irasnl  riiiii-tiniial  ai'tivify  ii 
siirh  slriirtiiiTs  as  niiisi-li's.  thi'ii'  is  a  \rry  iiiaikril  iiirrrasi'  in  liliioi|llii\' 
iliir  to  vasiiililatatiiiii,  whii-li.  in  part  at  li-ast,  is  ih-pi'inlriit  upon  tli< 
si'ri'i'tioli  into  till'  liloiiil  III'  siiiiii'  slllistanrrs  i'i'siiltill<;  I'l'iilil  tlir  Klallillllii' 
ai'tivitirs,  anil  is  not  I'litirrly  iliir  to  tin'  Mi-tion  of  \  asoililator  nrrvi'  liliris 

Similar  irsiilts  havf  Ihtm  olilaiiii'd  in  thr  rasr  of  thi'  ftiiiivriiis  wIhi' 
I'M'iti'il  to  sc<Trti'  li,\  till'  in|(ilinn  ol"  siTirtiii  ( si'i'  payr  4'J.''i).  I'nilri 
siii'li  riiinlilions.  thr  ox.xirrii  iiiiisimiption  has  Ihtii  oIisitvi'iI  to  iiicii'a^' 
alioiit   I'oiirl'olil  ami  to  In'  ai'ronipaiiir<l  li.\    ii   ililatation  of  tin'  );lanil. 

Till'  Work  on  till'  ki. IIII  II  has  Ih'i'ii  ispi'i'iall.\  intrifstinir,  lirrausi'  ii 
has  liri'ii  rminil  that  ini'ii'a>.i'c|  arti\ity,  wliirh  of  I'lmrsi'  is  iiirasiiri'il  h; 
till'  rati'  of  iirinr  cM't-rtinii.  is  not  alwaxs  arrompiinii'il  liy  iiii'ri'asr'l 
ronsiiiiiption  of  oxyiri'ii.  Wlii'ii  iliiiri'sis  is  prnilin-i'il  liy  inji'i'tini;  IJini; 
cr's  solution  into  thr  i-iiiMilation.  a  iriial  ini-n-asi-  in  iiriiii'  oiittlow  mav 
iMTUr  wilhoiil  any  I'haiijir  in  ox.\  (ji'ii  roiisiiiiiptioii :  whrrras.  on  tlio  otln' 
hainl,  wliiMi  »  iliiiri'tii"  siirli  as  soirMiiii  snlphalr  or  i-alTi'liii'  is  uscil,  tin 
ii\v'_'rn    I'lin^iiniption    inrrrasos    riiormonsly. 

[{•".'aiiliiiLT  till'  othrr  tissins  ami  iu-'.miis.  thr  (>.  I'linsiiniptioii  of  \\u 
liinys  aiiil  Iiraiii  appiars  to  Iw  smair.  It  is  a  \rry  siirnilii'aiit  I'art,  Imv^ 
('\rr,  that  till'  liii_'hrr  ri'iilnal  rniti'is  mit  i'\t  rniirly  si'iisitivi'  to  ilrpri 
vatioii   of  <>.. 

The  Blood.      In    the   lilonij    itsi'lf,  a   riTtain   amount    of  o\ii|atioii    yoi' 

oil   lu'raiiso  of  thr  pii'snii f  li'iii ylfs.     This  nxiilatiiui   lici-oiiirs  I'oi 

Kiili'i'ahli'  in   tlii'  hloml  of  animals  ri'inli'i-nl  aiu'iiiii*  l>y   thi'   iii.ji'i'tinn   >> 
phenyl  hyilrazin.     A  tlinrnnt^h   iii\  I'stiiration  of  the  eaiise  of  this  (jreatei 
(ixiilatioii   has   shown    it    to   he   owiinr.   not    to   an    iiiereasi"    in    niielcafi'if 
e'.irpuseles,    liut    to   the    pn-seiiee   of   the   yiiunf;   uniiueleated    red   bloml 


RKSIIIl     rilt\    IIKVONH   TlIK    I.I  N(iK 


397 


1  in|)iisi'lt'H,  wliii-li  ii|>|M'iir  ill  laiu'i'  imiiiiIm'in  in  lln'  I''      '  (imlrr  tlii'sr  cnii- 

hliiiiiH.     A  siiiiihir  iiificnsf  in  IiI'umI  uNiilaliim  <H-r  inu  poHlliciiior- 

iliMtfif  iiiu'iuiii,  till'  nilf  III"  iixiilalioii   ii inif  |>ariill<l  uilli   llif   lafi-  "f 

iiL'i'iKMiilinli  III'  I  III'  iril  ••iii'|)llsi'l('H. 


The  Mechanism  by  Which  the  Demands  of  the  Titiues  for 
Oxygen  Are  Met 

TImti-  air  1"«i  |in>Nilil.'  iihIIiimIs  l>y  uliicli  tills  may  ln'  liloliu'lit  almnt: 
•  I  I  li.\  a  <liani.'«'  in  llir  t'li  or  tlir  salini"  ridiHlitin'iits  or  tin-  lnii|>c'iiitnro  (if 
iIh'  |ilaNiiia.   so  tliat    tlir   ln'iiio;;|ol>iii   more   n-ailily   tli-livcis   up   its  loail 

nl'   (I.;   a 


III!  li.   I 


i\    an  incii'asc  in  tin-  mass 


iiiiiM'tiK-nt   III'  Itlooil  tliriiii);li 


till'  M'ssi'ls  of  till'  aftini;  lissmv 


I'l-i 


{{I'ttardiiii;  llir  lirsl  of  tlicsc  possiliililii's,  tlicri'  is  no  iloiilif  that  ncitls 
ic   proiliii'i-il   (liiriiii.'   liii'laliolism    of   ai-liii','   tissiirs.      As    we    liavi'   si-i'li, 

I  jii'ii  miisfli's  ( lrai-1    in   llic  prrsri f  an  almiiilMn f  O..  CO.   is 

iju I  ill  lai).'!'  amounts,  ami  wlini  llic>   coutrarl  in  a  di'tifii-iiry  of  (>., 

i-olai-tif  arid.      In  llir  sulima\illar.\    ylaml.  too.  it   lias  Im'cii  possililr  to 

w    lliat   till'  t'li  of  Ihr  \riioUs  l.loo.l,  as  iiirasuii'il  liy  tiir  \alur  of  A'  of 

f  li('iiioi.'loliin,  lirromrs  ilistiurtiv   iurrcasi'd  iliir- 


!lir  dissocial  ioii   rlirvr   o 

in','  '„'Iandular  arli\it.\.     Tliat  lliis  iiinrasr  in  ('„  will  dislodu'r  ( >..  wr  liavc 

ilirad.x    srrii    I  pajrc   :!>«i).      As   to   tlir    possililr   ililliirni f   local    rlialurrs 

III  triiii>i'raluic  and  in  salinr  roustilurnts  of  tin    plasma,  notliinii  ran  at 
I'lrsriit  111'  said. 

licuardiiu:  llir  srcond  possiliilily.  vasodilatation  iiia>  lir  drpriidriit 
•  illirr  upon  tlir  artion  on  tlir  lilood  vrssris  of  nrrvr  iiiipiilsrs  romin^' 
aioiiif  vasomotor  iiriv«'s.  or  upon  tlir  jirodurtioii   l(.\    tlir  artivr  tissue  of 

v.'isiNlilatint;   or    drprrssor    sulistanrrs    ( srr    paL'r    "JMt.        Much 


rviilriirr 


IS  I II   arriiitiulatini;  in 


1( lit    vrars    wllirli    triids    to   show     tliat    sllrli 


dr, 


iit'ssor  SI 


llistanrrs  arr  produrrd.  ami  tliry  may  hr  ritlirr  (1)   arids,  or 


i'l)  iir^anic  liasrs  of  a  similar  naluir  to  /Mmida/olylrlliylaiiiinr  (liista- 
1111111').  Tliis  lattrr  siilistanrc  is  of  ronsidrialilr  pli\  siojoirir  intrlrsf  lir- 
railst"  of  its  rlosr  rrlationslii)!  to  oiir  of  tlir  main  ainino  arids  of  tlic 
I'l-otriii  molrculr  liaiiirly,  liislidilir  ( M'c  pa^'r  t>(l4t.  its  I'flVrt  in  pro- 
duriii};  vasodilatation   is  i 


xtraordiiiai  \.     Tliiis.  lialf  a   milliiriimi  of  tlir 
II  lower  the  mean  arterial 


ilniu'  injeelrd  intravriiously  into  a  moiikrx    wi 
pirssiirr  liy  fifty  ]ier  cent. 

r.iit  before  siieli  an   liypotliesis  can  lie  entertained,  it   is  necessary  fn 
sliow  that,  intlepeiideiitly  of  nerve  impulses,  the  Id I  vessels  of  an  actinj; 


•  rL'an  liifiv  dilate.     The  liesi  evidenci 


lieeli  secured  liv  stildvini;  the 


rlTeets  of  stiniillatin!,'  with  eiiiiiephrine  t!i.    cervical  syiiipatlieti<-  nerve  to 
the  suliniaxillarv  ^land  of  a  cat.     The  Klaiiil  cells  iiei'ome  more  active, 


.    ! 


I.     fJ 


3f)8 


Tin;    UCSIMIIATION 


iiimI    (lil;it:iliim    of   tin'    .'irlriv    (irciiis.    iiltlioiitili    iiii    lilooil    vessels   nliiii' 
ei)iiie|iiiiiiic  In  similar  tlosaire  ])r(i<luees  ediistrietioii.     (>t'  enursc  in  sliciw 
ill};  tliat  liicil  eliemieal  |)riMliiets  of  aelivity  serve  as  the  exeitaiit  of  l(ie,i: 
•liiafatiiiii.  we  ijci  luit  iiwaii  to  imply  that  the  vasodilator  tiliers  jroiiiK  1" 
tlie  hlootl  \essels  are  ot'  no  use.     Indeed  \\v  know  tliat  Mieli  fil)ers  do  hr 
eoiiie  aetive  in  llie  ease  of  a  salivary  <rland  wlioso  eells  have  been  pani 
lyzed  li.\  atropine.  Init  it  is  a  si-jnifieant  faet  that  this  dilatation  is  of  rel.i 
tivel.v   shoil  ilnration,  wliereas  tliat  prodiieeii  hy  glandular  activity  last^ 
for  some  time.     The  siitrm'stioii  seems  theicl'ore  not  out  of  jilaec  that  uii 
der  normal  conditions  the  initial  dilatation  of  an  aetiiij;  <rland  may  ln' 
hroiiirlit  ahoiit  throiiuh  nervous  stimuli,  Init  the  later  dilatation  is  main 
tained  hy  iiietaliolic  jiroiluets. 


,  1  i 


CIIAI'TKK  XLVI 

TIIK  I'HYSIOLOOY  OF  BREATHIXC;   l.\  (O.Ml'RKSSKI)  AIR  AND 

IX  RAUEFlf:i)  AIR 

111  the  application  of  a  knowledjiP  "f  tlie  plivsiolo'iy  of  respiration  to 
the  investigation  of  disease,  a  group  of  conditions  arises  in  which  eon- 
sideriible  interference  with  physiologic  mechanisms  occurs,  not  as  a  result 
iif  disease,  Init  of  changes  in  the  atmospheric  environment.  The  regula- 
tion of  the  functions  of  respiration  depends  very  lartrely  on  changes  in 
the  physical  and  chemical  i)roperties  of  the  alveolar  air,  so  that  it  is  to 
lie  expected  that  similar  changes  in  the  atmosphere  will  have  a  marked 
iiitluenee  on  the  res])iratory  activity  .t.id  on  the  general  well-being  of 
the  animal. 

The  most  thoi'ou^riily  investigated  of  the.se  conditions  are  those  which 
develop  in  rarefied  anil  compn  -sed  air.  Either  condition  can  he  pro- 
duced exi)erinientally  in  the  laboratory  by  the  use  of  air-tight  chambers 
>|)iienmatic  cabinets')  and  suitable  pumps,  although  most  of  the  ini- 
imrtant  work  on  the  effects  of  rarefied  air  has  been  conducted  at  high 
altitudes,  where  the  l)arometric  pressure  is  low. 


MOUNTAIN  SICKNESS 

This  eondition  depends  jiriinarily  on  disturbances  in  the  control  of  the 
irspiratory  function,  and  it  is  on  account  of  the  useful  information  con- 
leniing  the  nature  of  these  functions,  rather  than  because  of  the  so-called 
disease  ilijclf,  that  so  much  attention  has  been  devoted  to  its  investiga- 
lin?i  dniinjjr  recent  years.  The  disturbances  iiroduced  by  the  I'aretied 
atiiiospheie  develop  rather  (luickly,  but  after  some  time  they  graduall,\' 
iiisapi)ear.  iiidicatint;  that  the  oi'ganisni  has  acclimated  itself — that  is, 
the  eonijieiisatory  meehaiiisins  have  come  into  play  to  bring  the  respira- 
tiiry  control  back  to  normal.  When  animals  are  placed  in  pneumatic 
liibinets  from  which  some  of  the  air  is  pumped  nut.  most  of  the  imme- 
diiite  syni))toiiis  observed  in  mountain  sickness  occur,  but  it  is  usually 
iiiil>raetieable  to  coiitinui'  the  observations  for  a  sufficient  length  of 
lime  to  allow  the  compensating  mechanisnis  to  develop. 

iiccause  of  their  great  value  in  revealing  the  nature  of  the  respiratory 
hormone,  many  of  the  results  of  the  recent  investigations  on  mountain 

.".99 


%  I 


4IM) 


Tin:   RKSI'IRATION 


!':» 


It 

:  ;  J  i 

liir 


sickness  liavf  Itcfii  {riven  elsewhere  in  lliis  volume  (  piijfe  :!(»()),  where  tli. 
j;einMal  syiniitoiiis  are  also  deserilied.     In  this  i)laet'  wc  sliall  considi  i 
very  liriedv  some  of  tlie  more  unitnil  iisimts  of  tin-  iiniililiini,  and.  nmii 
jiartiinlarly,  the  nature  of  the  adai)tation  that  oeeurs.     All  of  tlie  syiiip 
loms  are  essentialy  dependent  tipon  lack  of  oxyjjen.     Cyanosis  is  com 
iiion  aM<l  tlie  symptoms  are  iiiueli  the  same  as  those  of  cnal-^jas  ixiisonin;.' 
Xot  only  does  tliis  detieieney  of  oxy<ten  cause  acid  sul)stances  to  apj)e.ir 
in  the  liliMMJ,  thus  raisin;;  the  (',i  and  stiniulatiufi  the  resjyiratory  center. 
Imt  it  allows  other  poisonous  nniterials  to  apcuniulate.     These  act  on  tin 
various  nei-ve  centers,  ])rodnein{;  symi>tonis  which  vary  in  difTcrent  in 
dividuals  aceordin<^  to  theii-  relative  susceptihilities.     In  some,  the  difres 
live  centers  are  alTecled  and  nausea  and  vomitinj;  occur;  in  <ithors,  tin- 
hiirhei-  cerehral  centers  are  afTectcd,  causing  dej)ression  and  <teneral  nu'ii 
tal  a|)athy,  {.neat  drowsiiu-ss,  muscular  weakin'ss,  or  it  may  be  mentiil 
eNcitenn-nt  and  loss  of  self-control. 

The  susceptiliility  of  dilTerent  iinlividuals  also  varies  accordin<j  to  tlie 
nrnonnt  of  pi-evious  experienci-  in  nmunlaineerin;;  and  the  type  of  lii'eatli 
iui!.  Much  of  the  \alne  of  ))r(>vious  experiein-e  and  trainin;;  depeiuls  on 
the  aliility  to  perfoiiii  muscular  effort  economically;  to  ad.just  the  efToit 
to  the  available  oxy-ien  siip|>ly  without  i)ermittin{r  unoxidized  harmful 
l)roduets  to  accumulate  in  the  liody.  It  often  hapi)ens  that  lu)  symptoms 
a|)|>ear  so  lon{,'  as  the  i)erson  is  at  rest,  hut  immediately  do  so  whenever 
any  muscular  elTort  demands  a  much  nuire  alinndant  oxy{ren  supi)l.v. 

Tlie  typi'  of  hreathiii";  that  liest  witlistaiids  the  rarefied  air  is  slow  and 
deep,  rather  than  rapid  and  shallow.  The  reason  for  this  is  of  course 
that  miicli  more  of  the  outside  oxy{;en  {rets  into  the  alveoli  in  the  former 
case  fhan'iii  the  latter,  the  dead  sjiai-e  liein{r  iiractically  constant.  Tlw 
followinir  fi{rures  taken  from  observations  on  three  different  individuals 
will  illustrate  the  importance  of  this  factor. 


( 

.('.  I'K.ll 

NCI.  "►'  llfS- 

IIKKiliT  IN  MKTKl:s 

RKS 

■IKVriON' 

I'IKATIiPNS 
l-KR  MIM-TK 

AT  WHICH  .svMi'- 

TiiMS  OCCIiatKIl 

Siil.Jr.-t    1 

270 

'-•(1 

:•..■!  on 

i  t          t* 

tin 

It 

I'lOOO 

"     .-1 

70(1 

s 

f..'>nn 

( Fiiini   Iliilliliiirtciii.) 

After  livinsr  for  some  time  in  the  raretied  air  and  (|uite  independently 
of  trainin{r  in  the  ctlicient  performance  of  muscular  work,  adapltifitMi 
occurs,  so  that  the  symptoms  pass  ofT.  The  es.sential  feature  of  this  ndap 
tatioii  is  increased  absorption  of  (>,  into  the  blood.  Three  meehanisms 
liave  been  dcscriU'd  as  responsible  for  this  effect:  (I)  inerea.s<>  in  the  ten 
sion  of  t).  in  the  alveolar  nir;  (2)  assumptiiui  by  the  piiIiiKmary  epithelium 


m 


UUr.ATlMNd    IN    COMI'KIISSi:!)    AND    IN    UAI(KPIl:l>    AIH 


401 


(if  the  power  of  sL-crctin^  O^  into  llic  IiIihmI  :  ( :i  j  iiuTrasc  in  tlic  fryllirocyti'S 


111(1  licmo-rloliin  of  (li 


(1.    Tlh 


■I'ciisi'd  iilvcolar  O,  ti'iision  is  a  result 


pf  tlie  mure  rapid  lireathiiiir  l>riiui:iit  aliout  liy  tlie  iiu-reased  ("n  (if  the 
lidod.     11"  IK)  adaptation  (icciiircd.  tlie  ()_.  teiisinn  at  lO.OOO  feet  would  he 


ait  mill,  and  at   laJlOO  feet, 


i.S  mm.     Actual  ohservatioiis  mi  men,  liow- 


L'ave  at  lO.OdO  feet  a  tension  of  (>,")  mil 


d  at   l.').(»()()  feet,  o'J 


The  evidence   for  an   increased  seeretoi-y   activity   of  tlio   iiulnionary 
i'l)itheliuni  depends  on  ohservatioiis  made  hy   llaldane  and  his  eowork- 


wlio  found  that  hlood  collected  t'roiii  the  liiiircr  of  a  mail  li\  in<;  on 


hiu'l 


1   mountain    is   hriirhtlv    arterial,    whereas   if   this   same    hlood   is 


shaken  in  a  tlask  with  alv.-olar  air  from  the  man  frcuu  whom  it  Mas 
taken,  it  will  liecome  darkly  venous.  To  account  for  this  diiVercnee  it  i^i 
helieved  tliat  the  pulmonary  epitlulinm  forces  (>_.  into  the  lilood  contrary 
to  the  laws  of  dilTnsion. 

A  more  exact  ju'oof  was  sousrlit  for  hy  ciimparintr  the  reJaliNc  ;imounts 
of  <>.  and  CO  that  hlood  would  take  ii]!  ill  ulicn  exposed  diitside  the 
hodv  and  CI)  while  in  the  hlood  vessels.     Carhon  monoxide  h;is  a  very 


.'I'cat  aviditv  for  hemotrlohin,  so  that  ii' 


hlood 


is  sliiikeii  in  a 


Ihi 


ith 


air  eont.'iiniiifi  0.07  jier  cent  of  this  tras,  cojorimel  ric  measurement  will 
show  an  e(|nal  mixture  of  oxy-  and  c;irlioxy-heiiio'_'!oliin  Since  eail)on 
monoxide  is  destroyed  with  extreme  slowness  in  the  liody.  it  is  ))ossilile 
h'-  raiisin<;  a  man  to  hreathe  a  mixture  of  it   in  air  to  determine,  in  a 

•nio<ilohin   has  heen 


iox\ 


saiiijile  of  drawn  hlood,  whether  as  much  earl 

formed  as  In  rllro.     If  so.  tli(>  (V.  tension  in  the  hlood  must  ei|iial  that  in 

the  alveoli;  if  less  earlioxv-liemosrlohin  should  he  foriiied.  it  would  iiidi- 


ite  that  a  hiirher  tension  of  ()..  exists  in  the  hlood.     This  latt( 


the  n 


suit  whieli  llaldane  states  he  has  secured.  In  one  experiment,  for  ex- 
.iiiiple.  when  Idood  was  shaken  outside  tlic  hodv  with  O.dl  per  cent  I'O^, 
the  amount  of  earhoxy  hcmoirlohiii  formed  was  l!l  per  cent  of  the  whole 
liemotrlohin  ^Vllen  the  same  mixture  was  inhaled  for  three  or  four  hours 
the  jiereentatre  of  earhoxy  iK^nioiiloiiin  in  the  hlood  i-ose  only  to  "Jll  )tei' 
cent,  which  Would  correspond  to  an  o.  icnsioii  of  "_'■">  per  cent  of  an  atiiios- 
phere,  whereas  even  at  sea  level  the  tension  of  <*.  in  the  alveolar 
air  can  not  he  ahove  lo  per  cent  of  an  atmosphere. 


The  constant  low  tension  of  <  •_,  in  tin 
ipiiscles   and   the   perceiitaire   of  hem 


asiiia  stimulates  the 


d  h 


creiise( 


I  aft( 


...rial 
r  residence  for  s:ime  time  in  liii 


nil    til  liecome   marke(l 


III- 


iltitni 


At  lirst  this  is 


due  to  ii  eonceiitration  of  the  lilood  liv  a  dimiiiiition  in  jilasiiiii,  hut  irrad- 
iially  the  lilood-foiiiiin;;  oriraiis  hecomc  excited   and   an   actual    increase 


the  total  aiiiouiit  of  hemoy-lol 


>m  ocein-' 


!1  II 


li-rh.t  "f  t! 


use  facts  it 


is  interesllnur  to  eompaiv 


th 


era'jre  numoi'i-  ot    red   coriniscles   in   tlic 


hlood  of  inhahitants  livin<::  at  dilVerent  altitudes. 


ii        > 


4(lli 


Tin;    KKsmtATKIN 


'i::, 


I  I 

:  1 


H 


!    ,1 


m 


I 


:^||l 

i  1 1  1' 

1    i 

i 
j 

1 

■  ■ 

IIKI 

ilir  AIHIVK 

SKA 

li,KI 

1  COKriSCl.K.S 

(  MKITliS  ) 

( I'l^i: 

C.MM.  IlI.OOIl) 

fill  isti;llii:t 

II 

I.H"(1,<MMI 

Zlirirll 

111' 

."j.T.ll'.IMKI 

J>:i\t>s 

l.'illl) 

ti,r),"(l,(l(lll 

Ariisi 

1  sun 

T.ord.ddd 

('.iI.lill.TilS 

t.liU 

.<S.IM  1(1.1  II  III 

(  Fniiii  St:irliiit;. ) 

COMPRESSED-AIR  SICKNESS;  CAISSON  DISEASE; 
DIVER'S  PALSY 

Divois  and  i-aiNs.in  woikci-s  ai'c  Misccptililc  to  iii-ciiliar  syiui)t()nis. 
Tlicsc  arc  ficiiufiilly  nf  siitlicit'iit  scvcritx  to  i-aiise  death.  Iiiit  may  l)e  so 
mild  as  aliiiiist  to  escape  iiotiee.  The.\  lirst  ai)l>ear-,  iiol  when  the  worker 
is  suhjeeted  to  llie  iiiu'ii  i)ressuie.  Iiiit  after  lie  has  eome  hack  to  atmns 
liherie  iiressiiie.* 

AVliile  ill  tlie  eoiiipressed  ail'  the  \\iiikei'  as  a  rule  siilTers  no  diseom- 
t'ort.  A  stiitliiiess  may  lie  felt  in  the  ears  and  temiHirary  j-iddiiiess;  the 
resjiiration  and  pulse  rate  may  lieeome  slow  and  fre(|ueiiey  of  mietliritioi: 
may  lie  iiotii.'d,  hut  mine  of  the  symptoms  of  disease  ap|)ear  until  after 
the  eaissonier  or  <li\(r  has  heeii  deeompressed  (after  he  has  retuiiied  to 
atmospherie  pressure  i,  the  exact  time  of  their  onset  lieiii^  either  iiiime 
diately  aftei-  deciimpression  or  at  the  end  of  several  hours.  The  worker 
may  have  retiiined  lioiiie  and  spent  the  evenirifj  feeliiij^  perfeetly  well 
until  lie  went  to  lied,  when  symptoms  supervened  which  iiuiy  include  inus- 
<-ulai-  and  ,j<iiiit  pains,  verii-ro,  cmliarrassed  hi-eathiii},'.  siiheutaiieous  cm 
l)h.\sema  and  hemorrhaires.  jiaiiis  in  the  ears  and  deafness,  voinitiiijr. 
perhaps  hemopt.\sis  ami  epi-iastrie  jiaiii.     These  symptoms  usually  i)ass 

olT  aftei'  SI hours  liut  the  aithralfj;ia  and  niyali;ia  sometimes  i)ersist 

for  a  consideralile  time. 

In  the  mure  seven'  cases  the  lirst  symptom  is  severe  pain  in  the  mus 
eles  and  .joints,  ipiickly  followed  liy  motor  paralysis,  so  that  the  i)atient 
falls  and  is  likely  to  lieeome  unconscious.  The  juilse  is  almost  imper- 
ceiitilile.  the  it'spir.il ion  is  lalioreil.  sometimes  eve',  asphyxial,  the  face 
eyaiiosi'd,  and  the  Mil  face  of  the  liody  cold.  Many  of  the  eases  arc  fatal : 
indeed,  death  may  lie  almost  instantaneous.  Such  eases  are  eomnion  in 
careless  divinu'  ulieii  the  divers,  to  I'eturn  the  more  (piiekl.N-,  screw  u])  tlie 
outlet  valve  in  tlieir  li"lmels  so  as  to  (ill  their  suits  witli  air,  which  ear- 

•A  raiv..,..,  i.  3  Mvl  •  .  vv,.;,!.-::  th2!::br;  •■.-.•.•M  in  w::!.r  :,:,.',  ,.r,v.  ..t,  .1  fr..n.  ^!!i„s  l,y  ,,:,.atir  <.f 
c.imi.rtssfil  jir.  I'.ir  lli<-  |..•ls^.,^,■  „f  tin-  wirkiiicn  aii.l  iil  mau-nal.  iiili.  an.l  i.ut  nf  th.-  caisson,  ihi- 
latltr  IS  .ciiniitiil  Willi  .i  s,  ,  ,.n,|  sinalliT  i  hamlier  litlcil  Willi  air  links  :in,|  .li'<'.ini|irt'ssiti|i  c.jcks  A 
.iivrr  w.prks  in  a  wat.rj.r..,,!  .,,,1.  ij,,-  li,-a,|  lu-mu  nul.iM-il  in  a  mpprr  hi-lmit  Linncclcl  liy  hi.se  willi 
air  iiiini|.s.  l-.v.ry  In  inn,  is  ..r  .U  fr.-l  ..f  wal.r  <-.,rr.-s|„,n.|s  to  ,„»•  alni.ispli<Te  prt-nsurr  IIS  i.umiils 
lo  111.-  s.|nar,-  iiklil.  ...  thai  .il  this  ,|,|,ih  th.-  total  ,iir  |.r.  s-iir,  in  a  .aissciii.  .ir  in  :i  .livrr's  htlnift 
Woiil.l  ani.oml   lo  .In  |.oiiiic|s  lo  thi-   s,|n:in-   iikIi.  that  is.   i  1   .,lin..s|.hrr<'. 


IlUIATIIlVd    IN-    COMrRFSSKT)    ANP    IV    li AIllKlr.K    AlH 


403 


lies  ilit'iii  111  till'  surl'iicc,  vvlicrc  llify  ilccoiiiiHTss  tlii'iiisclvi's  li_\   iipcuiiii; 
the  valve. 

Aiiloimit s  (if  pci'soiis  (Ifiicl  (if  caissiiii  disease  liave  slinwii.  as  a  rule. 
intense  eiuiu'tstiiiii  nf  tlie  \  isceia,  lieiiKiriliajies  in  tlie  sjiiiial  eni'd  and 
I'laiii,  anil  eceliyinipses  on  llie  pleiiia  and  pei'ieaidiuni.  In  sinue  eases 
inferliiliai-  eniplix  si'nia  (if  the  liintis  and  laeeiatinn  (if  the  spinal  eni'd  and 
1  lain  liaxc  lieeii   initeil. 

The  Cause  of  the  Symptoms 

The  cause  f(ir  the  syniptums  is  nut,  as  was  at  (ine  time  suppiised,  that 
the  jiressufe  di-ives  the  IiIimkI  fmni  tlic  |ieiiplietal  iiitd  the  deep  fej^ions 
iif  the  iiddy,  inehidinu:  the  nerve  eentei-s.  Sueii  a  iii'dcess  is  iinpdssible, 
iiecause  the  tluids  (if  the  Ixidv  -and  all  tisstu's,  even  the  linnes,  ai-e  full 
nf  lliiid — are  inedinpi-essilile.  I'ressui-e  a]iplied  td  an\  jiart  of  the  body 
will  1)0  immediately  tlistfilnited  eipially  to  evei-y  dther  j)ai-t.  If  this  were 
nut  sd.  life  would  lie  impdssiiile  durin^r  nini  variatidii  df  atmospherie  pres- 
Mire.  It  is  now  clearly  estalilished  that  all  the  sym]it(iiiis  of  caissdii  disease 
are  due  to  (hrotiipn ssioii,  and  not,  in  the  sliuditest  dejiree,  to  the  meelian- 
nal  effect  of  the  |)ressure  itst'lf  i  I'aui  Heit.  Leonard  Hill  and  Maeleod  *V 

When  an  animal  is  under  jiressure,  its  tissue  lluids  d.ssolve  a  lai'iie 
MiMdunt  of  iras.  They  alisorh  it  in  oliedieiice  to  the  law  of  solution  of  a 
u'as  ill  a  lluid,  uliieli  states  that  the  aniount  nf  jri's  dissolved  in  water  is 
illi-ectly  pi'(i)i(irtional  to  tlie  ])artial  iiressure  of  that  <ias  in  the  atmos- 
liliere:  at  twd  atiiidsiiheric  )iressui'es  twice  as  iniicli  fias  will  jiass  into 
^iilutidu  as  at  /erd  ))ressure  (Dalldu's  law).  Sd  Idusi  as  the  pas  is  in 
-iniiile  sdlutidii.  it  dues  nut  in  any  way  ehaiiy:e  tiie  ])liysieal  condition  of 
'lie  lilood  and  tissue  fluids.  If,  liowi'M'r,  the  animal  is  suddenly  decdm- 
nressed  fi.  e.,  the  jiressure  (if  air  surroundiny;  it  is  reduced  to  zero).  tliP 
ilissdlved  iras  will  lie  so  (|uickly  thrown  out  df  sdlutiini  that  liulililes  of 
it  are  set  free.  These  linlililes  act  as  air  emiioli.  stickinn  in  the  puhiidiiie 
'■:i|iillaries  or  lildckin-r  uji  a  terminal  artery  in  the  liraiii:  nr  the>  may  lie 
■  iiu'e  and  tear  the  capillarv  wall  and  sd  lead  to  lieiiKirrhaire.  If  these 
Iiiilililes  are  pnidliced  in  the  pusleiidr  spinal  rodts,  intense  jiain  results; 
'i  in  the  anterior.  ludtdi-  paralysis.  P'rothintr  nf  the  lildoil  in  the  heart  im- 
["•(ics  the  actidU  (if  the  oriran  and  death  soon  fdlldws. 

The  fdlldwiii};  e\i)eriiiients  furnish  pindf  df  this  ex)ilaiia1idn:  A  frdir 
\as  placed  in  a  small  steel  chamlier  edlineeled  with  a  cylinder  of  cdiii- 
iii-essed  air  and  jirovided  with  twd  windows  liy  whieli  a  stronsr  are  liRlit 
'  "uld  he  passed  throutrh  the  chamlier.  The  ^cli  df  the  fddt  was  stretched 
"M  a  wire  and  fixed  sd  that  the  small  liiddd-vesseis  cduld  lie  seen  liy  apply- 
iiiir  a  mieroscopo  to  the  outside  of  the  window.  After  earefully  oliservinsf 
itie  circulation  of  the  lildod  in  the  vessels  at  atuidspherie  pressure,  n  posi- 


f 


■i  '; 


104 


Tin;  i{i:si'iiiATi((N' 


li\c  pf.'sMii,.,  .•iiii.iiiiitiri'.'  ill  s.iiiir  ('x|)t'riiii(>ii(.s  t(i  •  '>()  afiiiosplicirs,  \\,i. 
iiitiiM|ii,r,|  hut  Mo  (trc.l  .■(lul.l  lie  iii.tcl  (III  llii'  riicilliitili!;  1)1o<hI.  I'.n 
(ipfhiny  a  liqi  in  llir  cljaiiilicr.  (IccMiiii.i-fNsioii  to  zitk  prcssiirc  was  (luickK 

flVfrtcil   and,   iiii .lialci.v,    iaiLrr  lnil)l)ics   wviv   seen    to  dcvi-lop   in   tl,, 

''' !•  I'li"-I^iiiit  IIk'  v.'s>,.|n  and  pnidiK-in-,'  stasis.     Tlii'  luil.liN's  wfiv  dr 

lived  I'l-oin  till'  u'as  that  had  <:oii.'  into  solution  iiiidrr  ptcssurc.  On  ir 
aii|>l.viiii.'  tlir  picssiii-f  tlic  I.idibl.'s  of  jras  au'ain  went  into  solution  and 
Ihi'  Mood  ciicnlatcd  iioniiallv ,  When  the  pr.'ssiin-  was  siil)so(|ni'ntly  vci  \ 
iriaihially  hiwcnd  to  /cid.  the  ciriMilatioii  went  on  nndisturbt-d,  and  th. 
fro-r  was  rcniovi'd  from  the  ihanihci'  in  normal  i-ondilion. 

'''"'  I" '^^  involved  in  causiii!.'  I'aisson  disease  is  evidently  the  same  a- 

that   whieli  can   lie  ol.served   in  a  liottle  of  aerated  water;  if  tin rk   in 

siieh   a   liollle   is  drawn,   the  dissolved   j;as  eseajies  as   liillihles  and   elTei 
veseeiice  results;  if  the  liottle   is  recorked,  the  iras  reenters  solution  and 
the  iliiid  lieeoni.'s  ipiiet.     If  a  iiin  hole  is  made  in  the  eork.  the  <;as  \\'\V 
irradnally  I'seape  and  no  elVer\  eseenee  will   result. 

•  'onfirniatory  results  have  lieeii  seeured  hy  ol)ser\  ations  on  mammals 
'I'iie  arterial  Mood  jiressure  of  ralilnls  was  not  found  to  hecoun-  altered 
liy  exposure  to  compressed  air,  :iiid  various  animals  placed  in  a  lar;;!'. 
-troiMj;  steel  ehaialier  at  pressures  far  in  excess  of  those  to  wliicli  man 
e\.r  siil.jects  himself  di<l  not  show  any  symptoms  like  those  of  caisson 
sickness,  unless  the  pressure  was  siid.lenly  lowered.  Many  times  also,  ii 
symptoms  had  app.ared  they  could  lie  removed  liy  airain  sulijectiu';  the 
animals  to  the  compressed  air. 

IiivestiuMtioiis  wer.'  also  e.irried   out    to  determine  exactly  liow   much 

-■■!■>  ''"■  ''' I  of  an  jinimal  suli.jeele.l  to  hiirli  iiressures  coiitiiins,  and  liow 

lonir  it  takis  to  jilisorli  the  maximal  amount  of  L'as  and  to  release  it.     Ii 
was   found   that    the  lmscs  that    iiievejiscd   in   amount    were  nitro<ren   .■ 

ox.\u'eii.  and  that  these  lieeoi lissoheil  in  the  III 1  1 rdinjr  to  It: 

ton  "s  law. 

The  Prevention  of  the  Symptoms 

The    most    important    praclical   application    of    these    oliservations   con 
eeiMs  the   hiiL'th  of  tiiii.'  iv.piired   for  the  saturation  and   desatiiration  to 
-leciir.   for  tin    nsiills  sn-r<    <is  n  Inisls  uimn    irl,i,li   Ih,    ttafi-  ri<iiil,ition  „'' 

irnil.     ,„     ,n,. I, „;•<•:,, I    ,/„•    /,,,    „,.,,(    .-/.,    .'.,     ,  „,l:l  11,1 ,  ,1 .        Tile    lllosl     siu'll  ilica  II! 

outcome  of  the  alio\e  experiments  from  tids  stand|>oint  is  that  it  take- 
eonsidcialile  time  for  I  he  liloo.l  t.i  ;il,sorl)  its  full  ipiota  of  ;r;is  at  a  iriveli 
atmosplnric  piessinc  and  to  lilierate  it  au'ain  when  the  animal  is  decom 
presM-tl.     The  .••■uiM    of  d.  i.iy  i>  that  \\w  tissii.-  Ilnids  oilier  than  the  lilon.! 

take  miieh  lonjrer  than  would  1 xpecteil  to  reach  eipiililiriiini  with  tin 

partial   pr.'ssiue  of  .j.is  in   the  Mood   plasma. 


ai.ii 
la  I 


IIKl:ATIIIN(i    IN    ((IMI'KKSSKIi    AND    1  \    IIAl(l:Kli:l>    AIK 


40.1 


'I'll    iiiiilrotiiiiil    why    this  ili'hiv    slmiilil 


iicciir,    li't    us   Mi|i|i(is('   lliiit    til 


"iil.v    iriis  (•('filed  is  iiilnn.'''ii.     As  the  |iiissiirc  rises,  tlie  IpIhixI  in  the 

(■;i|iilliii'ics  (iT  the  liiiiu's  iiiiisl  dissolve  nil  luvreii  ill  |>r(iii(ntiiiii  to  tlie  pres- 
Miic  of  this  Hits  ill  the  iihcoli:  the  lilood  cjiiries  the  dissolved  <riis  to  the 
tissues  iind  these  dissohe  it  until  the  pi'essMlc  is  JiLjMill  eipmlizeil  lietuceil 
llielll  illld  the  lilooil.  'I'he  Mood,  aflei-  uivilli;  lip  its  cNcess  of  dissohcd 
iiilroireii.  retnins  to  the  liiiiirs  and  iisraiii  lieeoiiies  saturated  and  this  ^'oes 
le  ha\r'  lieeoiiie  satnraled  with  >j:;\s  at   llic  externa! 


until  Mood  and  tissue  I 
ssiire.   The    tissues 


\\\i 


third 


s   water   ami    tl 


ie\- 


iiitain    liii    iii;iii 


I'roiii    IT)  to  'JO 
tl 


It   oT  j'at.      I":jt,  1 


lowcvcr,  dissolves  li\('  times  more 
iseipieiiliy,   it   takes  loii'.'er  tor  a  iri^en 


iiilro'_'en  than  \\aler  I  N'eriioii  i 

Mihinie  of  tissue  Ihaii  of  Mood  to  lieeoiiie  saliirated  al   a   i;iven  incssiire. 


The   Moor]    in    mai iistiliites  one-twentieth   ol'  the   lioil 


\     wet 


rht 


that   il'  the  t 


issues  A\  ere  al 


Inpii 


I  tlh 


iild  d 


iss(dve  'H)  times  as  nine 


iiilrojreii  as  the  lilood,   Onaeeoiint  ol' the  fat  whieli  they  eontaiii,  1 
the  tissues  take  lip  more  than   tliis  jiroportion     iiamelv.   in 


ill!   averau'c 


iiinii  aliolit  :!."»  times  more  than  the  iiloi.d.     All  the  h 

:ilioiit  one  minute  to  eomplete  a  round  of  tl irciilation,  so 


ilood  III  the  liod\'  tal 


that 


III 


th 


IS 


time,  after  lieiin.'  suddenly  siilijeeted  to  an  i 

that  tlie  Mood  eireulates  oipially  tliPouirlioilt    the  liodv      the   t 


nereased  iiressiir(>     assiimiii'^ 


issues   \v  ll 


onr-tliirty  tiftli  satur;ite(| :  in  tl 


le  next    minute  another  tiiirtv-lifll 


thirty-four  thirty-fifths  will  lie  saturateil,  and  so 


Aft 


ll    ol 

er  five  minutes 


he  liody  will  1)0  aliout  '22  per  cent,  and   in  2'}  miiuites  alioiit  one-half, 
atiiraled;  liiit  //  will  tuhr  (ihoiit  tiro  hi)iir:<  hi  (nri   siitiinil 


lull    /.v   rilllll 


ilrh 


These  ealeiilations  assume  tluit  tlie  hlood  is  evenly  distrilnited  throiii.'li- 
Hiit    the  hody:   liut    this   is   not    the   case,    for    its   mass   movement    \aries 

e  mnsi'les 


loiisideraMy  in  dilTereni   parts,  lieiii-;  mindi  fireater  in  the  aetiv 
;iiid  in  the  trlands  than  in  passive  striietiires.  siieli  as  fat.     These  less 
i-iilar  parts  will  therefore  la;,'  lidiiiid  the  others  in  taUiiii.'  iijt  their 

ipiota   of  pas.   and   tlierefoi(>   iiroloiiu   tlie   time   n ssar.\     for   comiilete 

Mituratidii  of  tlie  body  as  a  whole. 

We  see  therefore  that,  after  some  time   i 


fill 


d   act 


11  compressed   air.  the  Mood 
ive  tissues   will   lie  saturated    and    contain    volumes   of  dissol\-ed 


'-'.IS  in  in-oportion  to  tlicir  relative  luilks:  the  fat.  aItliou<.di  not  saturated. 


yet    contain    ii])   to   fl\(>   times   more    eas   tliaii 


i|iial 


volume   Ol 


•  1.   illld   the   ])assive   tissues   will    he   incom))lptely    satuiati'd. 
esc  considerations  re<rardin<jr   the  sat 


Th 
if  the  body  .11ilil.\-   also   in    its  desat 


iiration   of  the   ( 


liir, 


rent    i»arts 


iiralioii.     Suppose,    t'or  example,   that 


the   external    ))ressure    is   suddeiil.\-    lowered;    the    blood. 
'iini.'s.  ^\ill  eoutnin  no  oxiTss  of  t;,i> 


leiivinjr   the 
leii   il    readies   the   tissues   it    will 
novo  <ras  until  the  pressure  is  ofpialized.  disehaiu'e  this  into  the  alveoli 


md  return  afrain  for  more.    Other  thinjis  boii 


e(|unl,  it  will  take  th 


it  tl 


I        ^ 


4()() 


TIIK    HKSI'IKATIDX 


W^ 


r-  1 

Hii 


<,:i 


sanif  iiiiinlitT  of  iiiinuti-s  to  dcMitiiriiti'  tliiit  it  took  to  sitturato.  niul  ih 
parts  lit'  tlio  liody  tliat  will  hii;  lu'liiml  tlic  otlicrs.  in  Ihmii^  dcsatiirati .; 
are  thoNo  witli  a  .slii!,'i.'isli  ciiiMihition. 

Wlu'ii  llic  mass  riKivfiiiciil  (.1"  tlic  Itlm.il  js  incifascil  In    iiiiisciilar  cmm 
else,  the  rati'  of  saturation  ami  ilcsatiiratinn  with  nitrom-n  is  inrri'asr  I 
in   proportion.      Diirin-j;  ai'tivi-   work   tin'   im-riasi'   in    nniviMiiiiit    of  tli 
lilooil  may  In-  four  or  live  tinn's  ovvr  tlw  normal,  so  that  thi'  tissni's  . 
the  raisson  worki-r  hm'omr  nnn-li  mori'  ipiirkly  ih'satnrati'il  ilnrin<r  ilfcom 
))ri'ssion  than  the  alioxc  lifinri's  would  li'ad  one  to  cxpi'i-f. 

Application  of  Foregoing  Laws  in  Practice 

Witli  regard  to  lliv  ajiplii-ation  of  these  i.rineiples  in  the  deeompressi..! 
of  eai.ssori  workers,  it  is  imi)raetieal)le  to  oeenpy  as  mueli  time  as  it  taN' 
to  saturate  tlie  hody  even  at  eoniparatively  low  jiressures.     Jf  the  giv^r 

dangers  attendinj:  work  in  i pressed  air  are  to  lie  avoided,  we  mu~- 

either  insist  on  very  •iradiial  deeonipression  or  we  must  show  how  ti 
dissolved  pises  nniy   he  ^rot   rid  of  hy  some  modilieation  in  the  deenii: 
pression  proeedure.     With  this  oh.jeet  in  \  iew,  we  nnist  determine  wli;r 
dilTerenee  of  pressure  may  he  allowed  lietween  the  external  air  and  tli- 
body  without  the  formation  of  huhhles.     Aetual  experienee  shows  tli:i' 
there  is  no  risk  of  l)idtl)le-formation,  however  ipiiek  the  deeonipressim: 
after  exposure  to  t   1')  pounds  pressure  (  i.e.,  2  atmospheres  ahsolute 
"Now,  the  volume  of  gas  eapalile  of  liein<;  lilierated  on  deeompressic.!. 
to  any  jriven  pressure  is  the  same,  if  the  relative  diminution  of  ])ressu!v 
is   tlie  same"— (Ilaldane    1.      On    reduetion    from   4   to   2   atmosplier.v 
the  .same  volume  of  gas  will  tend  to  he  lilierated  as  on  reduetion  from  -' 
to  1  atmospheres  -that  is  to  say,  no  liuhhles  will  form.     The  praeliiM 
eoiielusion  is  "that  the  ahsolute  air  pressure  ean  always  he  redueed  t- 
half  the  ahsolute  pressure  at   whii-h   the  tissues  arc  .saturated   withou 
risk."    Thus,  after  saturation  at  00  |ininiils  ahsoltite  pressure  (•  r>  afmos 
pheres"),  a   man  ean   lie   immediately  deeompressed   to  4."!   pounds   (      .' 
atmospheres)  in  a  few  minutes  wit' o^it  risk,  hut  from  this  i)oint  on  ijf 
deeonipression  must  he  eondueted  slowly,  so  as  to  insure  that  the  nitro^i  i 
l)ressure  in  the  tissues  is  never  more  than  twiee  the  air  pressure.     Tli- 
L'reat  advantasie  of  this  method  is  that  it  makes  the  greatest  possible  u- 
of  dilTerenee  of  pressure  between  tissues  and  blood  in  order  to  net  rid  ■■ 
the  tras  tli;>    'iiese  eontain. 

When  the  dci'ompression  f?-om  the  start  is  •rradnal.  the  desafuratin 
of  the  tissues  will  proirprssively  lair  behind  that  of  the  blood,  and  t!  ■ 
tendeney  to  the  liberation  of  free  gas  will  beeome  greater.  In  sueh  . 
ease  the  deeonipression  is  far  too  slow  at  first  and  far  too  rapid  latei 


nUKATIlINd    IN    ((IMI'KIISSi:!)    ANH    IN    UARKKIKK   AIH 


407 


Tlirort'f iciillv.  tht'i'tforc,  lln  thfamiin sxiini  xUoiiUI  In  nijiiil  til  /(cs/  ami 
rir;i  uluw  lalrr. 

Mcforr  rrciiiiiiMciiiliiii;  the  iiiloptiiin  (if  tliis  priiicipl"  of  stniro  <1<'. 
ciiiupfossidii  ill  caisson  \\(irU,  niildaiic  niid  Iiis  ('(iworkors  mailc  iiunirrous 
iiliscrvntimis  oii  llic  iiicitli'iicr  of  (Iccdinprcssioii  svniptnnis  in  lalmratory 
niiiiiials.  Tlu'.\  assert  that  tin-  sta^c  inctlKid  is  dci-iilrdly  safi-r  lliaii  tin' 
iinifiiriii  iiK'tliod,  tlic  advjintau'i'  Immiiji  iiarticiilarly  after  short  exposures. 
On  tlie  other  hand.  Leonard  Hill  eoiild  make  mit  no  ilefinite  advantau'e 
for  the  staue  liiethod.  The  t«o  methods  lia\e  also  heeii  compared  in 
actual  caisson  work  at  the  Klhe  Tunnel,  \\here  the  ]iressiire  was  :  2 
atliiosplieres.  Very  little  ad\anta<.'e  could  he  deiiioiistrated  for  the 
staifc  as  compared  with  the  uniform  metliod  at  this  coiiipaiativcly  hiw 
luessiire.  The  cenoral  conclusion  which  we  may  draw  is  that  the  static 
method  siiould  he  emi>loycd.  althoiiiili  it  is  not  to  he  e\|)ected  that  it 
will  alisoliitely  insure  alisenee  id"  dceoiii|)r<'ssion  syniptonis.  (if  cniirso 
the  (if'id  !i<lvanta<;e  of  the  sta-re  method  is  the  saviii'^  of  time,  makin<; 
it   ixissihle  to  persuade  the  workmen   to  adopt   it. 

There  are  two  other  factors  that  are  to  he  considered  in  liasteMiii<»  the 
de.saturation  of  the  tissues;  these  are  iiiiisi'iihir  r.nrclsr,  and  the  hrralli- 
iiifl  nf  an  huliljfrrnl  <ras. 

It  is  clear,  frcun  wliat  has  already  lieen  said,  tliat  tlie  <;as  dissolved  in 
the  tissues  \\\\\  lieconie  removed  in  iirojxirtioii  to  the  mass  movement 
of  the  hlood,  and  if  is  prohahly  true  that  muscular  exercise,  jierformed 
in  tlie  (h>com|)ression  chamher,  is  of  as  <;reat  importance  in  i)reventin!r 
the  sul)sei|iient  development  of  sym])tonis  as  a  much  prolonged  deeoni- 
pression.  Tn  a  man  at  rest,  the  circulation  tliroujih  the  central  nervous 
system  and  the  viscera  is  constantly  inllueiieed  hy  the  ))umpiii<;  action 
of  the  respiratory  nioxfiiieiits,  hut  in  the  capillaries  of  the  muscles, 
.joints,  fat,  etc..  this  infliience  is  not  felt  and  the  hlood  tlows  more  slowly. 
It  is  Ponse(|ueiitly  in  these  jiarts  that  hiibble  forniation  is  likely  to  oc- 
cur, especially  some  time  after  docomprossioii.  The  huhhles  cause  the 
neuralfiie  pains-  the  "lionds"  and  "screws"  so  well  known  to  caisson 
workers.  These  could  no  doiiht  he  entirely  ])re\eiited  h.v  jiuiscular 
exercise  and  ma.s.sajre  of  the  limlis  duriiijr  decompression.  In  illustration 
of  these  facts  the  followint;  exi>erinient  hy  (Ireenwood  may  he  cited: 
DiirinK  decompression  from  ■  "')  pounds  pressure  in  !tr»  minutes  "rirecii- 
wood  flexed  and  extended  all  the  limb  joints  .nt  frecjucnt  intervals,  with 
the  exception  of  the  knees.  Suhse(|uently  pain  aiul  stifTiicss  were  ex- 
perienced in  the  knees  and  nowhere  else."  In  another  experiment  the 
knees  also  were  flexed  nn<l  im  pain  was  felt. 

Rut  even  in  the  jiarts  with  active  circnlation.  the  iras  in  the  tissues 
may  lag  eonsidcrahly  behind  that  in  the  blood,  althuugh  the  dccompres- 


!    I 


.;:.-tt! 


I 

1 

i 


408 


KI.SIIKATION 


J    . 


**'""    'l"**    ' "    I'I"|>it1\     ,-..ll1lollc,|.       Thin    liilS    hiTII    sliinvii    l.y    la'OIUlf.l 

Ilill  III  tlic  .-.iM'  Ml'  till'  kiihi.'.v.     Tlic  "livMic"  ifiiH  ill   tliis  cum-  cim  1„. 
tiikcii  iiH  till-  iras  >li>sn|\..,l   ill  111.'  iiriii.'.  \t\    jiiiiilvziiii.'  wliirli,  tlicrcfor. . 
Ill   (lilVtri'iit   sliiifs  i>r  tlrci,iii|,i,'vsi.iii,   the  cM-css  of  iiid-oircii   „\vv   wIimI 
It   >li(.iii.|   Im'  at   till-  .•xtciiijil    jMi'sMin',   fiiii   l.f  ns.MTtiiiiicd.     (hi  dvvitiu 
Jtr.'>.si,,ii   l'r..iii      :!()  |«, till, Is  ],\    iu„  stac.'s  t..  /.t...  a  ci.nsi.iri'aMe  super 
satiiiatiidi  was  I'uuii.l  In  exist.     The  exeess  .if  nitmsieii  call,  litiwever.  Ii, 
eleareii  .mt  .if  lli.'  ki.ln.'.vs  lapiillv  aii.j  eoiiipl.tely  l.y  l.icat!  iiii;  oxyiren, 
vllieh   sli.iiil.l    III.  I..!-,, re   |„.   a.lliiiliist.'re.l   .jin'lii;;   .|.'e..iMp|-essi.i|l    ill   eas.s 

wliere  irn-at  .-aie  has  In  I..-  exeicise.l   I  I nard   Ilill  i. 

Whi'li   s\iii|>t(iiiis  il.i  a|i|iear.  they  can.   in    iimst    eases,   lie  relieved    li\ 

'■'■' 'Pi-.'ssi.iii.  ami  all  ni.iil.'iii  .-aissiiii  w.irks  are  i.r..\  id.-.l  with  a  special 

ehniiiii.'i'   f..r  this  puip.ise.      W..   iip,..|   s,-aieely  say   aiiylhin-.'  iilMiiit   this 
Ireiitiii.'iit   here,  as   its  \aliie   is  s.,  well   KiHiwii.     Siitliee  it   ti>  say   that. 
iiltlMMi!:h  it   is  iie.st   lik.'ly  t..  an'..r.!   i.'lief  uh,.,,  appli.'.l  as  s.m.ii  ns  pos 
sil.le  after  the  .'ippiaran..'  ..f  th.>  s\  in|.t.iMis.   y.-f   it    is  ..fteii  offieiu-ioiis 
when  applieil  several  .lays  after  tli.ir  ..nset. 

<iiiite  apart   frni  .  the  .laiijf.'is  .if  <|.'.-..nipressi.m.  it  iiiiist   of  e..iirsp  l..- 

reineinliereil  that  the  u.irkiiif:  < litions  in  a  caisson  are  s.uiiewhat  ilif 

fei'cnt  fr.iiM  tli.ise  at  atiii.ispli.Tic  pr.'ssnre,  as  the  air.  owinir  to  its  e..iii 
pressioM.  is  warmer  ami  is  loa.I.'.I  t.i  satnrati..n  point  with  nioistiire. 
This  h.  f.  M.'t  air  iMterlVr.'s  with  the  li.'at.reL'ulatinf;  mechanism  of  th.- 

ho.ly.  ixakiiifi  liar.I  museiilar  w.nk   very   un.M.mfortal.le  1 aiise  of  th.' 

temlem-y  .if  the  l..i.ly  temperature  t..  rise.  The  reaction  of  the  bo.ly 
iiiriiiiist  this  teii.l.'n..y  t.,  Iiyp.'i'th.'imia  c.msists  in  dilatation  of  the  sii- 
lierfi.i.-il  capillaries  ami   increaseil   heart   acti.m. 

When  siieh  w.irkinir  e<ini|iti.,i,s  are  re)>,'at.'.l  .lay  liy  .lay,  tho  appetite 

is  lik.'ly  to  fail,  i.aitly  1 ,nise  of  the  t.'n.len.-y  ..f  the  Ix.dy  t.>  suppress 

the  activity  of  the  metal.olie  in'.icess.'s.  so  as  f..  keep  .l..wn  heat  pro.liie- 

tion,  an.l  i)artly,  n..  d.iulit.  iM'.-aiis..  the  .lisrestiv,'  pr sses  nre  workin-; 

lieh.w  par  on  accuiit  of  there  l.eiii.^'  l.'ss  l.loo.l  .•ir.'ulalinir  thr.)UKh  the 
visceral  l.l.Hi.l  v.ss.'ls,  it  liavinjr  heen  sent  to  the  snrfa.-e  of  th.-  Ix.dy  t.i 
he  co..le,l  off.  Th.-  work.'r  theivfore  tends  to  take  l.'ss  f....d.  his  nietaho- 
lism  l.ecom.'s  .lepress.-.l.  ami  his  fact.irs  ..f  saf-ty  afjainst  bacterial 
infections  lie.'ome  lessened. 

The    risk    of  the   ai.p.'arnncr   .,f  sympt.iiiis   ..n    d npressi.in    is   also 

t'l-oatcr  when  the  air  in  the  caisson  has  heen  moist  and  hot.  f..r  the  lionrt 
has  heen  overworking  to  maintain  the  l.h.odtlow  in  the  dilaf.>d  vessels: 
it  trets  fatigued  and  is  consrqueiifly  unahle  to  maintain,  durinj;  deeom- 
pressi.m,  a  rate  ..f  lil<io.l(1ow  that  is  a.lc.piate  f..r  rarryinir  the  cas 
saturated  iilcod  to  the  lunjrs,  where  the  exce.ss  of  gas  beeonics  dissi- 
pated. 


RRI^:ATillN(t    rv    COMfRKSSKr)   ANh   IN    KAKKFIKh    AIK 


400 


Till"  crifcii-'ii  of  priipfr  Horkinir  i  •niilifiitiis  in   lln'  caiKKoii  is  tin 


liiif  the  wft-liiill)  fciiipcnitiiic.     Tliis  h1iiiiiI<1  s\t\i\i\  lifl 


OW      (i> 


I' 


inaiMtiiiii   tliix  riiii<lilii>ri    it    is   iM'crssnt'y    In    \  i>tililiil<>    tlit'   ciiiNMiiii,    |ii'cl'- 
cialily  wifli  iiir  tliiit   liiis  hccii  coiiIimI   Ii\  cDlil-wiiti'i-   ni<liiili>i's ;   in  nny 

i-lisc,    the    Vclltilntioil    sl|i>lll>l    In-    il(lci|llnti-    tn    ki'<|i    dnwii     tlir     wctiilllli 

IcinporHtiifc.      TIh'    iiifci'iisfd    i>x|>ciim'    of    vfiitiljition    with    i-iiiili'<l    nit- 
would  soon  \>f  liahiiHM'd  liy   flu'  tficatcr  woikiiin  tl1lcicm-y  of  tli,-   tiitn, 

IS  iissists 


CoiiNlHiit  circulation  of  'lie  iiir  in  flic  caissons  liv  means  of  fai 


Iso  ill  iiii|>id\  iiiif  the  funditions,  for  if  lu'lj)s  to  increase  dissipation  (if 
eaf   from  flic  body. 


ll^ 


w 


n\ 


I  ■  ' 


fi 


I' 


CIIAITKH  XIAII 

TIIK  Cim  I  |,.\T<»|{Y   AM)  UKSIMUAToUY  CIIANCiKS   At'Co.M 
I'ANYIMi  .MISCII.AU  i; X i: ur I SK» 

Dmiiiif  Hctivitv  tilt'  iinisclcs  iciiuiic  mmiv  tiiins  iimit>  liltiml  tlian  <lii 
iHK  rt'st.     Wli.'ii  the  arlivity  is  u  i.lcs|>icii<l  tin-  siicafiT  Itlmxl  supply  i 

pr.tN  i.lcil    liy    iniTi'aMMl   In-art   actiim   ac panii'il    liy   (lilatatiiui   of   lli 

iMiisciilar  aitiiiolcs  and  riiMstiictiiin  nl"  tlmso  nf  the  splanclinic  area.  -. 
fliaf  the  I'litiif  availalil.'  Iilixxi  supply  of  llic  luitly  is  iiiaiji'  to  cirrul;!! 
iiioiT  lapi.lly.     Wlifii,  on   thv  oIIut  liainl.  the  arlivity   is  .'oiilinfd  to 
liiiiitiMi  unuip  of  iiiuscjcs,  tlic  increased  Mood  sui)ply  is  mainly  pinvid. 

I>y  a  loeal  dilatation  of  the   hi 1   vessels  of  tin-  aetive  nt\iselcs  aeemi. 

panied  l.y  a   leeipioeal  eonstrietion  of  tliosc  of  inactive  jiails.      Tnd. 

these  conditions  there  may  therefore  l>e  no  (|uiekeiiinir  of  the  lil Kh. 

ns  a  whole.     In  order  that  this  accurate  adjustment  «(  Mood  supply  t. 
tissue   demands   may    he    promptly   und   adei|nately    hrounht    aliout.    ,ii 
iivailalile  types  nf  eoordinatini;  meohanism  are  called  into  jilay  ;  that   i- 
lo  sa\.  nu-ehaidcal.  nervous  and  liormone  factors  coo)terate  to  an  exten 
^^hich  is  dependent  upon  tlie  type  of  work  liein-r  performed. 

Hesides  the  ehantrcs  in  p;     .•  rate  and  hlood  pressure  which  are  cm 
denfly  desiijned  to  supply   more  Mood   to  the  acting'  muscles,   chanu" 
dependent  upon  a  secondary  elTeet  of  the  muscular  nnivements  have  al- 
to he  considered.     Althousfh  the  various  factors  work  together  and  ai' 
more  or  less  interdependent,  the  tiiud  effect  can  he  iniderstood  oidy  aft. 
we  liave  studied  the  relative  inMueiiee  of  each  separately. 

The  Mechanical  Factor.^   It  is  parti.Milarly  with  rogard  to  this  fact., 
that   the  circulatory   clian(;es   may   l)e   an   unavoi<lalile  conse(|uonce   oi 
rather  than  a  useful  adjustment  to.  the  muscular  cfTort.    The  cITects  vai 
with  the  type  of  exercise  performed.     In  repeatedly  liftinR  and  iowerin 
dinuMiclIs  from  the  tloor  to  ahove  the  head,  the  cniitractinsr  mu.seles  . 
the  hack  anil  extremities  and  of  the  ahdomen  eomi»ress  the  veins  an 
cause  the  hlood  to  How  more  rapidly  into  the  heart,  so  that  the  arlori; 
pressure  suddeidy  rises.     So  lonsf  as  this  compression  exists,  the  veil 
remain    relatively   empty   ami    the   .Trteries  overfilled,    hut    whenever    r 
ee.ises  .^nd  the  muscles  relax,  the  veins  fdl  up  auain  and  the  arterial  prr 

the    circulatory    anci    rcsi'iratory    ailjustmcnls. 

410 


«ll,\N<ll:s    AIKIMI-.WYIS'II    Misn  l,,\K    I  M  K<  IK| 


411 


.urt'  iiiarkfdiv  falU,  until  llic  I'xtra  ^pHcr  in  the  xcinn  Iihh  ln'<'n  m-ciipii-il 
ly  IiIikhI.     If  iw  I'm-  thin  miKon  tliiit  llw  iirti'riiil  hi I  prcKNtirt'  is  iilwavs 


foiinil  to  Id-  little,  if  aM\.  nliov*'  normal  ulirn  taken  willii 


n  a  U'w  st'i'oiiils 


ifti>r  Hiich 


CXl'I'lMSC, 


II 


.sulisi'i|iii-ntlv    iisi's 


fU 


III 


'caw^c    till'    ollii-r    fartors 


i'i'N|iiinKilili>  for  till'  iiii-iTa'*i'i|  jireHMiir  ii|iiii'k  lirart  aiid  artrrioli'  I'luistrii'- 
lioii)  ail'  still  ill  ii|ii'ration  at  tin-  linn'  tin-  \i'iii>.  aiiaiii  lin-onii'  fillnl  with 
Mooil.     Till'  pnri'ly  itiri-haiiii-al  inlliirni-i'  outlasts  tlii"  rsriiMsr  fur  a  roin- 

jiaralivi'lv  short   linn',  \\hrn>as  th-  iu'imium  ami  lioniionr  iiilliu'i s  nm- 

i'li|iorti'<l   liy  till'  olisi'i'vation   that 
1    wht'M   'tic  siiliji'i't   is  li't't  stamliiii; 


tiiiui'  ai-tiiit;. 


Tl 


us  inti'i'|)i'i'ta* 


till'  fall  iif  hlooii   pi'i'sMiri' 
aftrr  a  trivi'li  aiiiouiit  of  ilir 


I'll  111 


liiwril  to  sit 


.ith  li 


is  I'lhows  ii'stllit:  oi 


ill  tl 


sure  oil  tlu'  ahiloiiiiiial  ■     n 


caiisi's   iiiiii'i'   lilooi 


il    t. 


■.s  :i  lit 


I'WIS      I. 


H 


I'iiij;  puii'l 


fiillou  iii^  iluiiilihi'll  I'  'i^<  I'  i 
timis  art'  iiiailc  at  i-lo.s,  <  h  i/i 
Till'  liiri-liaiiiral  frspoih  ■  i  i 
ilii'oinfli  till'  rati' of  fillinir  of  ti  ■. 
in  a  hraltliy  romlitimi.  it  will  i. 


'u\t  position  till'  pri's- 
itii'  I'lTi'i't  of  gravity 
r    •    ii'i'i-         iilloii,    Itappiii't   ami 
K     M  -ril      ,,  till'  pri'liiiiinarv  fall 
I)     .ii'iii.i  .tiiiti'il  if  till'  ol)si'f\a- 


t   I'Mi   i'  (       !()   I'xi'rrisi'   arts   llii'ri'fori' 

!'  Ii<!ii     .  n!   M I,  aiiil  if  this  ori;aii  is 

tlir  itiT  intliiw  liy  I'orri'spoiiil- 
iiijrly  inrii'asi'il  ilisi-liai-KC  Liki-  i'mtx  oIIiit  |)liysiolot'ii'  iiii'i-liaiiisni.  tin- 
heart  works  with  a  larne  faetor  of  sjifet.\  a  reseivc  power-— ami  it  is 
the  rate  of  venous  (illiiifr  that  iletermines  how  iiiueli  of  this  I'eserve  must 
he  ealleil  upon  to  maintain  the  eiieiilatioii.  In  isolatei!  liearf-liinir  pre))- 
arations  Starling  ami  his  eoworkers  have  very  elearly  ilenioiistiateil  tl..' 

I'lose  ilepeiiileui f  eanliae  output    Upon   rate  of  venous  lillinir  ami  the 

eiiornious  raiiire  throiiirh  \\liieli  the  systolie  ilisehar^re  ean  he  iiiaile  to 
vary  by  alteriim  this  faetor.  As  explailieil  elsewhere,  when  the  reserve 
power  of  the  heart  is  lesseiieil,  the  rise  in  lilnoil  pressure  followiiicr  cxer- 
ciso  is  liinirer  in  attaining  its  uiaxiniuin.  \\liieh  is  set  at  a  hifrher  le\el  ami 
persists  for  a  longer  tinu".  Ohservatioii  of  the  extent  of  these  ehannes 
furnishes  a  most  useful  flllietiolial  test  of  eanliae  e(1lieieiie\ . 


Other  meehnnieal  faetors  that  auirineiit  the  eanli 


thf 


reaseil 


resjiiratory   iiioM'uieiits.     Diiii 


itpiit  ilepeuil  on 
>ii'atiiiii   the   in- 


■ase  in  eapaeity  in  the  thorax  eauses  hotli  jiii  opeiiiii>,   ii]i  of  the  thin- 
lleil  veins,  so  that   hloml   is  asjiirateil  towanls  them   from   the  extra- 

oii  of  the  hlooil  \essels  of  the  luiiirs. 


thoraeie  venous  system,  anil  a  ililatati 

so  that   the  hlooil  (imls  its  way   from   riirlit    to  left   heart 

Although  this  dilatation  will  at  first  tend  to  eause  more  lilood  to  eolleet 


more  reai 


lil> 


tlu-  ititrathora 


eie  vessels  ;ui( 


d  less  to  111 


puniprd  out  i.f  tin  in,  ihe  expira- 


tory net  when   it   supervenes  will,   liy  eomiiressiii!. 


tl 


le   veins,   eause 


thr 


extra  blood  to  be  cxjielled   into  the   left    ventriele  and   tlienee  into   the 


!  -i        ft 


f  'i 

It 


vi« 


412 


Tin:  Ki:si'iitATii)\ 


MIL' 
I'    ill 


ilft.Tii's.      Il   is  nl.vioiis  tllill   ilM-lv;i.sf.l  ,|,.|)lli  :iii,|  |'r,..|i|..Mrv  ..I'  tli.-  |fs|.iii, 

Iniy   iii<iv.iii,.|iiN  will  : |,.,iit.'  Ihr  lil.>...lll.,\v  .-iikI  tcii.l  to  Pilisc  llic  iiiL 

I'iiil  111 1  picsMirc. 

The  .-ilioM.  )';irl..rs  will  mi'  inln  play  .lining'  mnst   kin. Is  i.f  iniisciiliii 

f\ricis.-  siii'h   as   walkiiiu-.    nimiiiii.'.  or  suiiiuiiijr  .hmilil.rlls,  <.tc.      TIiit. 

iiiv  .Tilaiii  t\|H's  of  s,-uiai-  .iVoil,   liowcv.r.  in   w  liicji   ih,.  iiif.-liaiiir,i 

fiirlois  i.nMJii.-,'  .I.'ci.lrclly  ,/).s/ , /;•/./»-/  ,//,,/.v  oil  tin-  i-iiviilat ion.  Dm- 
il  siisiaiiifil  rlTori  as.  lor  ,.\ai,i|.l,..  in  piilliin;  auMinsI  a  r.sislan.-..  ,. 
.•itli'inpliii-  to  lift  a  lira\y  loa.l.  tlif  r.-spir,itioiis  arc  siisp,.|i.l.Ml.  oftni  aflri 
.■I  <I<'.'|)  irispiratjnii.  ami  the  .'ontiartnl  alM|,,ininal  iiius.-lcs  press  the  ,|i,-, 
I'lir.-iuin  lip  into  the  thoraeie  eavity.     After  a   preliminary  sipiee/iiiir  out 

"''  ''' '   '''■-•  "'■  ■ill    l'l-"'ii   the   veins  of  the  ah.loiiien   into   the  thorax   aihi 

then  iVoin  those  of  the  latter  into  the  systeniie  aiteri.'s,  with  a  eonse.pieni 

rise  in  arterial  pr.'ssine.  then mes  to  I.e  a  .laiiiinini:  hiiek  of  Moo,}  into 

tlie  jieripheral  \eiiis.  eallsinjr  them  to  swell  an.l.  if  eontiniieil.  inarke.l 
.■.vanosis  may  .levelop.     \V||,.n  siieli  elTorts  are  maintaine<l   for  Ion-    tin 

.•iHerial   pressure  he-ins  to  fall,  ami  this   fall  is  very   pr in 1  imi | 

ill  the  elnl  of  the  .iTort.  iMe.iuse,  the  eoiiipressioii  heili!:  leinoveil  from  tli. 
.•ih.|oinin;i|  ami  thoraeie  x.ins.  th.'se  open  up  iuul  form  a  larjie  iinlill.ij 
hloiiil  reservoir. 

•^   ^i'liil'-if  II haiiism   (Mimes   into   play  .liirin-:  ex|)ulsi\e   aets  such   as 

ilcfccation.  i.artnrition,  <te.      In  these  the  -loltis  is  close.l.  usually  aft' r 
n  preliniin;iry   inspiration,  ami   a   pouerfii!   exiiiratory  movement  "is  pel 
forme.l.  with  the  eonsc,,uence  that  the  ill t rat hmacie  am!  intraalMlomimi! 
IiresMires   rise   e,,iisi,|eralily.    -reatiy    auemeiiiini,'    the  systolic    dischar-i 
.•iiiil  eansim.r  ,i„,   i,|,„„i   ,„,,..„,„  ,„  ,i„,       |.,,,..|„^,>  „,•  t,",,.  „i,sf ,.,„.,i„„  „, 

""■  ''I I'l""    '"   t'"'   l:iii;''   veins  of  the  al..|omen   and   thorax,   however. 

the  later  effect  of  the  elVort    is  to  diminish  the  sy.stolic  discliarv'c.  hut   till 

fall  in  liluoil  pressure  which  this  would  I xpcctc.i  to  occasion  is  nmskcd 

The  pressure  reimiins  hiLrl,  | aiise  other  la<-tors  iiicrcasinj;  the  i)eriphciMl 

'■''^'^'••' ''""I"  ill!"  |il;i.\-     The  fall  ill  hlood  inessiirc  followini.'  these  act. 

miiy  I.e  very   marked   iiuhed.     Similar  mechanical   ctTccts  are  produced 

in   the   .-lets  of  coliehiiiL'.  sl zillir.   etc. 

The  capacity  ,,f  the  veins  varies  coiisideralil y  with  the  position  of  tin 
iiody.  and   it  is  in  oii|er  th.it    we  tiia.v  c;iuse  jiltcrations  in  this  capacitv 

nnd  therefore  encouia-c  ,i  moie  r;ipid  M Illow  tliat  w  <•  st  retell  tile  I.od> 

.'ifter  sillinu'  for  soiii..  time  in  a  ciamped  iiosition. 

The  Nervous  Factor.     The  v.i-us.  vas nstrictor  ami  respiratory  cen 

ters  ar.'   ;dl    eN,-i|,.,|   ,|iiiin.^-   muscul;ir  etTort.      In    the   cjirlier  stML'cs   tli. 
cNcitatioii  depends  eiitiielv    on  iicrvoiis  impulses  transiiiittc.i  |o  the  ecu 

ters.  li:it  later  il  depends  on  chanu-es  in  tl m))osition  and  temperatuiv 

of  the   Mood   ilowintr   throu<;h   tiicin-tlu.   lioriiume    ffU'tof.     Tiic   initial 


lal 

1 
I' 

1    1 

mL,. 

<II.iS(ll:S    ACIO.MI'ANVINC    MISCI'LAK    KXIKCISi; 


41.1 


stiiiHiljitidii  oi'  tlic  ccnlcrs  iiiust  l)r  line  lo  ccirln-jil  impulses  iiiiicpiMi<l- 
ciitlv  triiiiMiiiltcd  to  tlic  iilxivf  cciiti'is.  >iiiiM'  the  (luickciiiiijr  of  tlie  pulse 
;iii(l  respirations  iiiiiy  lie  ojiseivcii  to  lieirin  liefmi'  the  aeliial  inuseuiai' 
eoiiti-actions. 

The  Hormone  Factor.  ^Ve  have  to  eonsidei-  first  tlie  nature  of  the 
horiMoue,  and  si ndly  the  modi'  of  its  aetion. 

The  Xatinr  of  tin-  I/oniuims.  The  most  important  lionnone  is  car- 
lionic  aeid,  l)Ut  when  tlie  exercise  is  strenuous  and  eontinued,  or  from 
tlie  very  start  is  of  sueh  a  natuie  that  it  uses  up  oxyireii  more  (iiiiekiy 
than  the  blood  can  supply  it  to  tlu-  museles,  laetie  aeid  also  appears. 
Kvideiiee  for  these  statements  can  I'cadily  lie  supplied  in  man  liy  analy- 
sis of  the  expired  air  (for  earlion  dioxide)  and  of  the  urine  I  for  laetie 
aeiiH  before  and  duriui:  museular  work.  The  real  hormone  in  both  cases 
is  believed  to  be  an  increase  in  the  llion  concent  ration  of  the  blood. 
There  is.  howevei',  no  diiect  ])ioof  of  this  assertion  that  is  to  say,  no 
one  lias  actually  shown  that  a  measurable  ehaiiire  in  the  Il-iou  concentra- 
tion of  the  arterial  blood  (  for'  i<\'  course  a  cliauire  in  the  venous  blood 
>\ould  be  of  no  sis;nilieance  >  docs  occur  before  the  chaniies  believed  to 
be  dependent  upon  acid  prodiietioii  make  their  appe.-ir.inee.  The  well- 
known  bulTer  aetion  of  the  blood  (that  is,  its  ability  to  take  up  eon- 
siclcrable  ipianlities  of  acid  or  of  alkali  before  any  iiereeptible  ehaiiRe 

occui's    in    ll-ion    i 'cntrationi    fur'iiishes    another    reason    why    doubt 

must  be  cast  upon  the  II  ion  hypothesis.  The  most  delicate  means  for 
deirjonstratin^  a  chanue  in  M  ion  concentration  of  the  blood  consists  in 
lindinu'  the  dissociation  constant  for  hemo-ilobin  ami  the  results  have 
shown  that  acidosis  develops  durinir  exercise  at  least  at  liiirli  altitudes 
'  HarcrofI '  i.    So  far  as  w  i'  are  aw  are,  how  ever,  it  has  not  been  ])ossible  by 

direct    mc-isuieinent    (  pa  ire  '_'!!  i    to  detect    a    rise   in    ||-ii>ii   coi nti'ation. 

<M"  course   it    may    wrW   be   that    the   si  nsitivcucss   of   the   various    nervi- 

centcis   and   other   strnetui'cs   towards   the    ll-i oMciiitration    is   \  er\ 

iinu'li  ui  cater  than  our  most  retined  and  si'usitive  Ijdioratory  methods 
can  i,\eal.  Such  is  at  least  commonly  believed  to  be  the  ease  for  the 
respiratory  center  (sec  p,ii;-e  :;.")!  ).  and  it  uia\  ,ibo  be  so  for  those  .if 
\aseulai'  tone  and  cardiac  action.  It  is  ne\ crllieless  pnssible  that  an 
increase  in  liie  free  carbonic  ;iiid  itself  the  earbon.itc  anion  i  JICO  i. 
in  other  Words  is  the  elTecti\e  iioinioio'.  Ill  the  lii'st  sia'.'cs  111'  muscular 
vvork.  this  ini'i-case  \miuI'I  be  due  lo  'greater  production  of  ('<».,  whereas 
l;iter.  espei'ialiy  when  tlo-  w.ok  is  sircinuius.  lactic  acid  wmild  dceum 
l>ose  the  XalK'O  of  tiic  Mdod.  jiber.il  inu-  IICU.  which  would  becom.' 
.'iddcd  to  that  still  liciiiii'  |irocluci'd  liy  the  aelixe  muscji's,  ani|  jis  the 
X.ilK'O,  iliulTer  substance)  bccauo'  irradually  used  up.  wnnhl  cause  a 
rciati\ely   friiatci'  and   irrealcr  iiroportion   of     ll('(»     to  exist    in   a    free 


i! 


f'i! 


:y 


"M 


'i 
1 


h,! 


V, 


*>  i 
1 


'isl  M 


414 


Tin:    KI.SI-IRATKiS 


stiitf.  TliHt  the  CO,  tension  .,)  thi'  alvfoljn  air  >li(>nl<l  |p.-  I'i.uikI  In  li. 
louiTi'd  I>v  iin.lonu'ctl  niusculiir  cxcrciso  in  md  way  detracts  from  tin 
oxpIaiiatioM,  for  if  is  (lepenilent  upon  the  jrrcatly  incrcascl  rate  .. 
Iiioveiiieiil    „(  air   into  .ind   ont    o|    the   alv.'oli    i  see   also   |.a^'e    -Wii). 

One  serions  (lirtieultv  in  aeeeptiny;  tlie  Il('<»,  ion  as  tlie  exeitiiiK  lior 

' "'  "•"  '111'  iii'rve  ( ters  dnrinir  inuseular  exereise  depends  on  tlie  ol, 

servatioii  that  tlie  alveolar  CO,  afte-  some  time  is  lower  than  normal 
If  we  aeeept  llaldane's  teaeliini.'  tlial  there  is  aeeurate  eorrespondone, 
Ix'twfen  the  tensions  of  C(t,  in  arterial  Mood  anri  alveolar  air  net  oiil> 
dnrin-r  rest  l.iit  also  durinir  mns<'iilar  activity,  then  ohviously  wo  iinist 
discard  the.  IICO  hypothesis.  Le.Mnird  Hill  and  Flack.  •  however.  ha\. 
shown  ipiite  .jearly  hoth  in  experiiiu'iital  animals  and  in  man  that  eoui 
liliriiim  helween  the  lilood  and  alveolar  tensions  of  CO.  may  fail  t.i 
oi'cnr.  AVIien  l.lood  with  I'xcess  of  CO,  is  iiije.ted  into  the  .jtl«illar  v  ,i 
of  doj.'s.  the  respiratory  eeiiter  is  stimulated,  as  shown  liy  the  ineroas«.i 
hreafhini:.  which  indicate.,  that  the  CO,-rieh  blood  must  have  passe,! 
thron^rh  the  Iniiu's  \vithont  the  excess  of  CO,  heini;  removed  from  it 
Hill  heli.'ves  that  the  .liirusioii  of  CO,  out  of  the  l.lood  into  the  alveolar 
air  may  lie  depressed  in  niiiseuiar  exercise,  and  that  this  rather  than  tti.' 

appearanc'-  ,,f  lacti,-  acid   iii   the  |,| |   is  responsihie   for  the  low  CO,  ten 

sioiis  nsiially  toniid  present  i  se.-  pa^'e  :!(HI).  He  points  ,.nt  in  support 
of  this  \i,'«  that  a  person  after  evercise  ean  hohl  his  lireath  for  a  iTiiieli 
shorter  time  than  is  iisu.il.  and  the  CO.  meanwhile  mounts  in  the  alveolar 
air   \er\     rapidly 

The  only  «ay  l.y  which  process  imiy  lie  made  in  a  prohlem  like  thai 
ntider  diseiission    is.   however,   to   adopt    some    hypothesis   and   tlien    to 
Karli.-r  evidence   for  or  airainst    it.     .\t   the  pr.'sent   stai;e  of  our  kiiowl 
ed-fc.   th.'   hypotlirsis   usually   adopted    is   that   a   sli!,'ht   ehansro   in   Il-ion 

'■'" ntration  oi   the  M |   is  the  etTectual   hormone.     It   is  an  hypothe 

sis  which  is  snpp.irted  hy  the  parallelism  Let  ween  the  effects  observed 
diirintr  niuseiilar  e\..reise  an<l  those  produced  hy  exi>erimental  increase 

ill    ll-li'l nce?iti;it  ion. 

Thf  nil,, -Is  „i  th,  Ihinintiii  These  may  he  classified  as  follows:  (1 
strictly  lueal  etTects  on  the  iiMiseles  themselves:  I '_>  I  elTeels  on  till'  heart 
and  ■:;  etVc-ts  on  ih,>  ncrv,'  centers  Tji.'  local  prodnction  of  acids  n 
till'  miis,-|es  uill  cause  dilatation  of  the  arterioles,  for  it  has  l.eeu  shown 
hy  \arious  ohsciv.'is  that  acids  cause  relaxation  of  vascular  mtisel,. 
Mvcn   the  capillaries  themselves  are  said   to  he  dilated   l.y  earl.onic  aci,l 

(Sc\erinii      Tl iVccts  proiiu I  on  the  heart  hy  chanires  in  ll-ion  eon 

eentration  of  the  l.loo,|  have  t.eeii  particularly  studied  l.y  Starliii;.'  am! 
F'attcrsoti,  '  who.  \\(.rkin-.'  on  isolat.^d  heart  liinir  )nepnralions.  hax. 
shown   that   the   luMit    relaxes  more  .-md   mor.'  and  .liseharu'cs  less  l.loo.i 


if 


(IIANCiKS    A(  r(IMPA\YI\(!     Ml  SCII.AK    F.XKRClSr. 


415 


;iN  tlif  II  i«iii  (•uiicfiitnitioii  of  the  pcrliisioii  lluid  is  iiiciciiscd  l>y  addiii}? 
(•()_.  to  thi'  ;iir  vciitiliiliiii;  tlic  luii^s. 

Tlif  iiitl  iciicf  iif  cliaiifju's  ill  11  imi  (•.luccntiatioii  (tf  the  Idood  (in  the 
\aj?ii.s  and  \a:^|'nlot^>^  cciitfrs  is  usually  l)i'lii'\fd  to  he  stiinulatoi'v. 
Thcri'  is  no  doulii  'liat  an  i'lcrcasi'  in  ('h  suinulati-s  tin-  vasoconstrictor 
(inters,  not  only  of  ilic  iiicdulla,  Init  al>o,  although  much  more  fccMy. 
..I'  tlic  spinal  cor.l.  Hut  it  is  a  i|Ucstioii  wiictlicr  any  part  of  tlic  rise  in 
s\stolic  picNsuic  diiriiii;  muscular  exercise  can  be  attriiiuted  to  this 
<ausc,  for  the  encpiniously  increased  liloodllow  >vhicli  is  known  to  occur 
makes  it  |)rol)leiiial ical  whether  any  \asoco)istriction  leally  oci-urs.  If 
It  does  so,  it  must  lie  conliiicd  to  the  splanchnic  area,  where  it  would 
have  the  elTect  of  lirmnin!.'  alioul  a  reciisi  riiiution  of  the  total  available 
Mood  hy  expressiiij;  it  from  the  vis.'cra  and  seiidiiiH;  it  to  the  active 
iiiuscies. 

The  elTeet  of  increased  11  ion  concentration  on  the  vastus  center  must 
l>e  insiiiniticaiit.  It  is  coiiimonl\  hclievcd  'hat  it  would  cause  not  what 
;>  actually  oliservcil,  a  (|uickeniin.'.  liul  rather  a  slowini;  of  the  lieai't  rate. 
Hut  even  this  is  doi.  itful.  The  slowinir  of  the  heart  tliat  is  oliserved  in 
ispiiyxia.    lor   cxamjilc.    is   in    part    at    least   due   to   the    increased    intra- 

ciaiiial   pressure,   for  when   the  carotid   artery   is  ci cted   wilh   i!   mer- 

rury  valve  so  that  the  blood  escapes  as  the  pressure  rises  abine  the 
hoiiiial  le\cl.  iMi  slowing  of  the  heart  is  saiil  to  occui'  in  aNphyxia.  As 
Leonard  Hill  and  Klack  ■  have  sliow  n,  however,  a  part  of  the  slow  nit:  is 
lue  to  the  diri'ct  .•tTect  of  <'<•_..  If  increase  in  the  ll-ion  cMincentratio-i 
Iocs  alTcct  (he  heart  diiriiiir  muscular  exercise,  it  must  act  li>  inhibitniu' 
the  vairus  tone,  which  is  opposite  tci  the  action  which  it  is  usuallv  be- 
lieved lo  have.  The  activity  of  the  respirai..ry  center  is  of  course  e\ 
riti'(l  by  increase  in  ll-ion  concentration,  and  this,  as  we  have  seen,  will 
laiisi-  important  i-ImIIL'cs  h  the  rii-cubilloli  because  of  the  mechanical 
■iTects  whicii    follow. 

.Monir   with   hormones   we   must   eonsider  the   ciTeet    of   ehantre    in    the 

■  1111)1  nitiin    iif   till    hhiiiil.      That    this   rises   dnrinir   muscular   exercise    is 

.11  known,  but   that   it   should   be   responsible    for  many  of  the  cardio- 

aseiilar  ad.iuslmcnts  that   oecui-  is  ipiiti'  eonimonly  overlooked.      It    is, 

lor  example,    very    likely    that    rise   in    blood   temperature   is   responsible 

H    the  aceeleralion  of  the  lieart  that   occurs  duriiiu'  excfcis.'  when   both 

,e_'i   liave   bei'ii   severed,   anil    it    no   doubl    is    responsible    for   a    |)art    ,it 

.  ist  of  the  vasodilatation  and   lesjiiratorv    ac'clcration 

I'iiiallv.  it   is  iiiteiestiiii:  to  speculate  as  to  the  nature  of  the  ehaiiKes 

Itat  occur  when  the  "second  wind"  is  aciinii'cd  during'  strenuous  miis- 

ilar  exercise      In  ruiiiiinii,  U<v  example,  eonsiderablv    more  distiess  is 


^pe|•iell !    a   short    lime   after   the   start    th.'iii   some   time   later 


Th 


^lli 


4iti 


llli:    HI  SIMHMION 


vcrv  (Ifliiiitc  ilijinjxrs  occur  at  tlic  time  llic  irlid"  is  cxpoiiciicccl-  iiamil\. 
a  sluwiiit:  iiiiil  stcii.lviii(.  of  the  iMcvioiisIy  iimcli  (luickoiiwl  and  iiTi'du 
lar  i)iils.'.  Nwciitiiis.',  aii.l  a  marked  fall  in  the  iT.spinitoty  (|U«ti('iit.  Tl- 
last  inciitioncd  chaiiirc  j.ossihly  jrives  a  cliic  to  tho  cause  of  the  otlici- 
111  the  early  stas;es  //.  <f.  is  raised,  which  indicates  that  relatively  nior. 
CO,  is  beiiij,'  expeil.-.l  from  the  Mood  into  tho  alveolar  air  than  oxy;;.  u 

IS   hein^'  ahsorhed.   iierliajis    heeause   of   iiia«le(|uate   niovenieiit   of   1.1 ! 

throu-rh  the  Junsis.  At  the  time  of  the  adjustment  it  is  possible  that  ,, 
pronounee<l  vasodilatation  o.-curs  in  the  mnscles  and  coronary  arlcrii- 
The  former  ehan-;e  hy  loweriiifr  the  arterial  Mood  pressure  will  relie\. 
the  pumpin-r  action  of  the  heart,  and  the  latter  will  improve  its  power  m 
contraction  liy  siiiiplyiusr  it  with  more  oxy^ren. 


m 

i,., 

ii 

i 

i 

1^ 


u 


KKSIMUATIOX   HKFKK'KMKS 

( Mono!,'raphs( 

"•"'■"']];,;'■■      ''■'"•   •'■'•"l-i'-atorv    r .,,„„   ,.,•   ,|„.   |,|.„„|_    |„i^,,r^i,j.   ,.„„,_   ,',„„|,H,|., 

H..rr„l,o,.   H.:    \:,;,.,.| '.   ||,o„|l,„,|,   ,|,.r  l'hv.,..l,.«i,..   I-m,:,    i     m, 

""'"    M''i,''   niu  '•'*■.,'/,;■«"''"'""  'I''-  A""i,nir  l..im   McLH.h.n,   Kr«,-l,Mi Irr   l'l..v,i.,; 

"'"'     Loi"i'I^?  I!.l^'''"'"    '*'''""■"-•    '">'n,;.li.,„:.l    M,.,|i,,.l    M.,„„,n.plM.    K.    A,„„|.! 

K.itl,,  Arlloirr     Tl„.  M,..  l.nim.n  .,r   l;,.s|,ir:.ti„M   in    Mn„.  l-„ril„.r  A,K,o ,  i„  IM.v- 

<ili';,'.v,  K.  AmimI.I,  l.nnildn,  liidll. 
>^.l.,M.k,   y.:      li,M,.|vali(.n  ,1.  r   Alnuiii-,  Kr-.-Lni-M.  ,|,r   l'liv.i„|„«i,.,    Ijios,  p.  d.-,. 

(Oriifiiial   Articlesi 

iK.ith.    Vrtllur:     Ct.   K'lirlhiT  A.hiiiu,- 
ll.".^.r.  .-.  K.:     And.  Int.  M,.!..  |.p|  ;.  x,i,  -i,;  ,|,i,|.,  |.,|;    ^^    ;„, 

.;",';         '':■  '""■".""■'■•    ^-    '•■•  "•"'    ^'■■l""y.   'I.   K.:    Aim.  .I,„m.    l-|,v.inl..    Iltir,   ,)     |,, 
'\Iill/,r.  S.  .1.:     .Iniir,  Ihysii.l.,  Isni',  xiji, -Is. 

Il.il.l.oi..,  .1.  s.,  loi.l   l-ri.sil.v.  .1.  (1.:     .|,',„r.    I'l.vsiMl..    llMi:,,  xwii    ■•■..-, 

Il.il.hoi.    .-omI   llniiirl.-i*:     Jl.i.l.,   l!l|:!,  xK,  :;:;.-,  •     -   ■ 

•ll.n.|,rs„n,    V..   r|,i||i„i.„  „rt  |,    :iii,l    Wl.i'tM.v':    Am.    .loiir.    I'hvsini.      Is.l,-,     vxxviii      < 

ll.'ri.|.i>..n  nil. I   M„rn:<-.:      .|..ur.   l!i,.l.  Ch.  i,i.,    \\i\- ,  sw,  ji; 

"Kn.-li.  A.  .-111.1   l,ii„||,i,r.|:     .loin.  I'ln  »i,.|..   Il.j.l.   xl^i.  ;;ii-  il.i.l      im:    li    -,.| 

MV:.r.-..    I;.  (;.:     Am.  .1.  ui    l'l,v«i„l..  Mil:,  xliii,  7;;;  ,|,i,|..  i!ii7.  xliv   :;i;;. ' 

^•M.'lMM.k     l;.:      Sk;,ii.l.    \,.l,.   I     PI,yM..I..   11.11,   XXV,   s::   .■„rf,.r.   K.   W: ,.    Kx,.. 

.Mf.j..    li'M,    X  X,   L'  I . 

"  I'.';.1m.,Iv,  I'.  W.,  ;,,i,|  W.nlw.Til,,  .1.  A.:     Ai.li.  Int.  .\!...l.,  11.17    xv     11,! 

iil.iiMs.  T.:     ■Iimr.  I'lu^i..!,.  liHis,  xxxiv,  L'l:;    ■'■..■; 

i-f..rt.T,  \V,  T,:     .l.mr,   I'li.vsi.il.,   Is'...",,  x\ii.    l.V,. 

'  <liiiMi;iii«.ii  ;iii,l   ||;,|,|t,ii,-,  .!.:     .I..iir.  I't.v.-K.l,   lull     xKiii     '7' 

"li..,.ll,l.y,    W.    M ,,  ,H,.I    lt.iry,    1'.    |i.:       Am.    , r.    I'liv,!.,!',.    1.7|-.     xxxvii      Cl     -,1 

H<...llil.y.  W,   \|,,  :,ii,|  SlLMm.,ir,  V    \.:      Il.i.j.,  p.   lis.  '        '  '    ' 

'   ■^l''"'*.  N-   •'■•  I  Si anil..!,:      .hmt,   Physii.l,,   lim.-).  xxmI,  ,;ii, 

'■•Suit,  K.  II,:     .l.iiir.  I"li\,«ii.l,.  llMis,  xxxxii,  ::ii| 

':.si.«:irt.  (i,  N„:oi.|   I'ik,.,  P,   II.:     ,l„„r.  nivM,',!,.   I1..17,  xx    r,| 

i-.Cnoml,s.  II,  <•,,  „„.|   I'ik...  K    II,:      |'„„..  s.mv   Kx|„r    ISi„l,  .M,„|      li,|s    x>     .V, 

•'Kh.uI,.    A.:      Sk.-.ml.     \  ,,1,     (     Pi,v.l„|..    1.,,,,.    xxiii,   J|s;    ..„„|     \.    Ix„.nl,'  «ii|.    .\|,,, 

Ixi.iuii.  ii.i.l,.  17:., 


CUANOKS    .\l(<)MI'ANYlX(i    MlSt'lLAK    KXKKflSK 


417 


•Uulilaiio,  .1.  ^.,  uml  I'ricNtl.y,  .1.  «i. 


.Idiir.  I'hvsidl.,  I'.'li'i,  xxxii,  UJ.".. 


- "Seott,  K.  \V 


Am. 


Jcpur.  I'liv.xinl.,  1U17,  xliv,  liMi. 


Ni-wl.iiru',  MiiiiiH,  Jinil  I'orlir,  \V.  T.  :   .'our.  V.\\h-t.  MimI.,  liilrt,  xxiv,  ". 


K\. 


--llassolUilili,  K.   A.,  iiiiil   l-Miiils).'!'!"''!.  '"'"■■ 

Skaiid.  Ar<-li.  f.  Physiol.,  ll'l:;,  xxmi,  1:i 


iHookiT,  1).  I!.,  Wilson 


I).  \V.,  and  Conn.tt,  H. 


-i(ani|.l).-ll,  .1.  -M.  M.,  UouKlas,  f. 


.1  llob-ion, 


'l.inilharil,  .1. 


.1(1 


Ihiil.,  I'.ti:'..  xivi. 


riiysiol.,  I'.lll.  \xx\  lii, 


Itioili.ni.  Ztsihr.,    I'.H:;,  xxx\iii,    ..,  ai"! 

Am.  .Tour.  I'hysiol.,  li»17,  xliii,  :!.'i7. 
1".  (i.:     .lour.  Physiol.,  I'.IU,  xlviii, 

llaldanc,  .r.  S.,  and  Douglas,  (".  O.: 


Dou^jlas,   ''.   li 


.\rt,    Kryi'lmisM' 


diT    Phvsiiiliiyif 


.M. 


uiiyrii|il 


:il«a 


>fl.  .1.:     .vo    K<s|.iriiti.ry    hiiurtioni 


Itlood. 


"Milrov,  T.  H.:     (.^uarl.  .lour.  I'hysio 
I'Kh-t.-l'ier,  \V.  M., 


1..   I1M;{,  vi,  :iT 


I   ll"|ikins. 


V.  »; 


Jour.  PhvBiol..   l!"i: 


;t7  ;  also   Kl»^i-hfr, 


\V.  .\l.:     .lour    Plnsi.d.,  I'.M.l,  xlvii.  :!l!l 


■'i;v(Trl,  .T.  II.:     Pro 


I'hVM.i 


.rour.  I'hvsiol.,  I'.iu".'.  sxxix. 


ilVml.r<'V.  M.  S..  and   Alliii.   R.  W.:      .lour.    l'!i>«i. 


d..  I'.Hio. 


■hurkmasiir,  (• 


,l<i 


Ph\> 


id.,  i!ii:,  li.  lo.-). 


■Uiiujjla.i.  I"    (i..    llaldanc, 
lioy.  So.-..    l!'l\   -K": 
^'11  ill,    l.conari 


S.,   ll<'iid«'r><oii. 


id   Sidini'i 


1- 


l.T.   i:.   !• 


i.    Ma.-l.'M.l.   .1.    .1     K. 
Oroenwooil.  M..  Kluik,  M..  I'ti' 


.    Physiid..    I'.iii:;.    xxix,    .'i 
Hill 's  ("aiKS.m  .Siokiicss. 


Ilalda 


.1. 


I  In 


Wat. I     Itiiia;;. 


CcillllllittlT     of      till' 


Adii 


Phil.    Tr 
ihll.    Lroll 


I'.lltisli 


Hill  s  CaiMson  Sirinc 
•;Cotton.  T.   K.,  Ha(i|i"rt,  and   l.ouis. 


■  Hill,  I..-onard.  an 


,1  .Mail I,  .1.  .i:k. 


ll.'ail.  r.ii>*. 
.lour,    Phv»i(d.,   liloS,  xxxvii. 


■"Patterson 


S.  W.,  1- 


ir,  11..  and  Starlmy 


K.  II. 


.lour.  Phvsiol.,  I!il4,  xlv  iii,  4ti; 


y 


i 


Mi 


PART  V 
l)l(;KSTI()iN 

CIIAPTKR  xiAiir 
CKNKUAL  I'llYSKiLOdY  ((I'TIIK  DKIKSTIVK  CaiANDS 

Tlic   I'liiii-iioii  of  ilicrsthiii   is  to  liriiitr  the   food   iiifo  s<icli  a  coiulilion 
that  it  .-an  l.r  iiitsoilpr.l  tlin.iijfii  llic  iiitcstiiiiil  t'pitlicliiiiii  into  the  l)lo..,! 
iiiiil  iMiipli,     ('ail)nli\<iiiitcs  arc  lin.kcn  down  as  far  as  iiionosai-cliaridcN 
iiiMilral    fats  arc  spiil    into   fattv   a.-i.ls  aii<i   «:lv<-<Tiiu';  and   proteins  an 
l.n.|<.M    clown    into   the   amino   acids.      The   au't'iicics    uliicli    clTcct    tlicv. 

ilcconipositioMs  arc  tlic  digestive  cn/.ynics,  or  ferments,  eoulai I  in  tin 

various  di-.'cstivc  llnids  or  juices.  Tlie  diircstive  juices  are  produced  \>\ 
■.ilands,  wlii.'li  are  most  numerous  in  the  upper  levels  of  tiie  nastr.. 
intestinal  tract,  the  lower  levels  havinfr  as  their  main  function  that  oi 
al.snrption  of  the  diu'csted  prodm-ts.  In  onler  that  the  masses  of  fon.l 
may  he  kept  in  a  state  ..f  proper  consistency,  and  that  they  may  nnn, 
readily  aloni;  tin-  di-icstive  canal,  numerous  mucous  ".'lands  are  als.. 
>c;itt.  .d  alon-;  the  wlioic  extent  of  the  canal.  Some  <d"  the  di«estix. 
•flaiMf  such  as  the  main  sidivary  ■rhinds,  the  paiu-reas,  ami  the  livei 
'li-^i'  c  llieir  sccieijoMs  into  the  digestive  canal  liy  speci;d  duel- 
-       iithcrs,  v:i.-h  as  the  isolated  salivary  u'land  follicles  in  the  moutl. 

•  j..>lric  jrlands  an.l  the  crypts  of  Lich.'rkiihn  in  the  intestine,  do  (e. 
Ii.  ■  an  .in.-itomii  ally  diNtinct  duel,  Imt  dis<'har';e  thi-ir  s.'cretions  din-clh 
int. I  the  iliirotive  tnlie. 

It  will  lie  convenient  t nside?-.  first  cd'all,  certain  properties  i|i:it  ai. 

'•'"" 'I    '"   '•>'•   diircsti\e   ^dands.   .ind    then,    the   conditions    under    wlii.l 

I'.iili  L'hiiicl   finiciiiinatcs  durini;  dij;estion. 

MCROSCOPIC  CHANGES  DURING  ACTIVITY 

Sirncinrally  the  .Mctixe  pait   (d'  the  ;:lands.   rcprcscnti.l  l.y  t\w  acinii 

'"■  liii'i'le.  IS. iposcd  of  a  l)ascment  mendiraiie  liiu'd  internally  with  tin 

secrelini.'  e|,itl,elinm.     (Mitside  the  l.asal  mcmhraiu'  are  the  lymph  spa.r 
an.l  I.I.hmI  eapillari.s.     Aft.^r  the  y^lai',.1  has  1 n  at   rest,  the  cells  l.eeoiii: 


|-HYKIOI,0<IV    OK   TIIK   KKIISTIVI'.  (il.ANOS 


410 


(illo,l  with  Knu.i.lfs  or  smnll  jjlol.ul.s,  wl.iH,  an-  oftn.  so  iM.inrnms  .is 

„1 St  .•ntir.-lv  tn  ..l.lit..rat,.  llir  MU.I.us.     Wl.rn  tl.r  i-'la.ul  iMnmn-s  nH.v. 

„„  „,..  ,„h,r  ha.Ml.  tl...  trranul.s  or  «1oIm.I.-s  Lav.-  Ilo-  .rlls.  ,.x.M.,.t  lor  a 
f,.«    «ln.-li    nMnai.i    louanl    11..'    luin.-n    l.or.l.r.      ( I'lu's.    14:!   an.)    144.) 


I'lK     11.1.      I 


,n..,l.i.il.-    -..r.l ;    (.    iiH.i    l'r.,1 1    -■••l '•'■■"'    i-'""^'' 


Thoso  oLscrvalions  in.lirat.-  tliat  the  trraiiular  or  -loLular  .natrnal  must 

rcprcsHit  part  at   Irast  of  tl...  s.MMrlioi.  of  11..-  Klah-is.     S •Iiiiu-s.  .'vt-ii 

l,,.for.-  Il.fv  arc  .•Mru.lc.l,  th.'  -raimirs  lu inr  .•liaii-nl  into  some  .lifl.-r- 

,.,.t  malrrial,  as  is  in.li.-al.-.l  l-.v  tl...  fart  thai  thry  stain  .lilT.rrntly  from 


A 


rf\ 


W^J 


\  ■^- 


^'4.5A' 


^ 


--^V-c   ^^^^1 


♦^ 


J  » 


I   * 


I 


r,^.    u,,      l..„.„„l    .1...,.!    ..f    r.l.l.,.    ...    sa,>l..v    -.,,..-   ■■(    ..-m-v    .^..m.n,,l    „,    fr.-h    ■-.a..^      Tl..; 

"...-' M.I....-I -■■  ■—  „,'■;-. ■,:^:; ';„;-' :;:;  ,;::;' ,:;;: , ,:  :j!;!  L:::T%l 

.l.ll>.IV     liV      111).  >I. lit      111...    ..CI..!  .      .....I      tl.l       I'A..     ......      ...1... 

|..ih.iif   ...TV.-.     I. Mil.    l,....Kl.y.. 

thosr  of  thi'  rrstinir  u'hiM.i.     It   imist   not   ho  th.-ii-ht,  h..w('vtM-.  that  an 
.slriisioii    of    ^'raniih^s    „,  r.  ^sorlh,    afconipaiiifs    s.cirtory    activity,    for 

un.h'r  .••■itaii i.litions  a  ...pi-iis  so-n'tioi;  of  water  aii.l  i.i..r«ani.'  salts. 

as  well  as  a  .■tM-faiii  ai.H.iint  of  or-aiiic  material,  may  he  pi...ht.-e.l  with- 


:i| 


420 


t)ioi:sTio\ 


out  any  vhnuuv  in  tlii-  jii  i!iii.„'rni.iit  „(  Uw  Kiiimihs.     In  .siidi  cases  it  li;i~ 
l.foii   ol.scrv.Hl,  as   in    fli,.   pancivaH,   tliat    tin,-   .•liiiiincls  d.-vrlop    in   lli. 

|i|-(il(.|.|;i,|||    ,,|'    (||,.   ,.,,|1    I  see    |.,|ir,.   4-J!)  i . 

Fidiu   this    liistoli.-ri,.   cvidfi it    w.nild   appear   that   the   K'laiiil   cill 

•liiriMi:  rest  is  fn.li.w.Ml  uith  llir  :.r..p.rty  of  huihlinn  up  out  of  fh.-  pro 
toplasiM.  as  uraniil.'s  or  (.'lohul.s,  thr  material  nliirli  is  to  s.tv."  as  one  oi 
tlie  main  (.iijaiiie  eonstituents  of  the  seeiction.  It  is  eoramonly  Itelieve.l 
that  this  is  the  preeiitsor  of  the  aetive  fernieiit  of  the  scoretion;  henee  ils 
name,  :!i,iiof,,ii.  It  lias  been  shown  that  the  process  (.f  sej)arntic)n  of  tiir 
zymoiren  irrannh^s  Marts  around  the  nucleus  with  the  |)roducfion  of  a 
l)as(,pliile  suiistanee,  which  in  hardened  specimens  sometimes  takes  the 
form  of  tilaments.  From  tliis  basophilic  >  ni,itfni,l,ism.  as  it  is  called,  the 
granules  .ir..  jri-iidnallN  formed,  and  then  for  some  time  continin-  to 
inider-o  sli^'ht  further  chan-es,  as  is  evidenccl  hy  the  fact  that  th.' 
stainini:  reaction  of  thos<.  near  the  l.ase  of  the  cells  dilTers  from  that  of 
those  at  the  free  manjin.  When  the  trland  cell  is  excitd  to  seeretc. 
the  (rniiniles  I.efore  l.ein-r  extruded,  as  noted  al.ove,  oft.'ii  nn<lerK<i  a 
definite  chanire,  1 miiiu'  swollen  and  mine  ulohular  in  shai>e. 


MECHANISM  OF  SECRETION 

These  hislolo-ric  studies  merely  tell  us  that  active  changes,  associate! 
with  the  i>roduction  an.l  liberation  of  certain  of  the  cimstituents  of  its 
secretion,  are  .pccurrinf;  in  the  t:land  cell.  I.uf  they  throw  no  light  on  th. 
mechn.dsm  wlu^rehy  th."  s;lan.l  cells  s,.cretc  wat,  r  and  monjanic  ,ulls. 
This  may  he  d.'iun.l.nl,  to  a  certain  extent  at  least,  on  dilTerences  in 

osmotic   pressuiv    ,.,T   p;,;r,.    II,,     A   possil Nplan.ition   of  the  tlow   „f 

water  is  as  follows:  If  a  watery  solution  of  som.'  osmoliealjy  active  siih 
stance  is  i.ut  in  a  tube,  which  is  closed  at  one  en.l  by  "a  mt'iiibraiie 
inipermeabl..  to  this  substance  ami  at  the  other  by  one  i)erineable  to  it 
and  the  tube  immerse.l  in  water,  a  continuous  current  will  b.. 
found  to  issue  from  the  iMrmeable  end  so  lonsf  .is  there  remains  am 
osmotically  active  substance  in  the  tube.  If  we  assume,  then,  that  the 
membranes  at  the  two  ends  of  the  secretiinr  ceil  are  of  such  a  nature  that 
the  <.iie  next  the  basement  membrane  is  impcrme.-ible  to  some  osmoticallv 
active  substance  mannfactun'.l   by   the   cell,  .ind    the   other  toward   the 

lumen    is    i.ernieable,    it    will    .|..,.,r    that,    so    long    as    this    substance 

exists  in  the  cell,  it  will  attract  water  from  the  blood,  an.l  the  wat.r 
together  with  the  osm..tically  active  substance  will  be  discharged  into 
the  lumen. 

It  is  possible  that  when  anything  eycites  the  cell  to  secretory  activity, 
such  as  a  nerve  impul.se  .>:    li.irm..ne.  it  does  so  by  causing  a  change  in 


rilVflol.iMlV    UF    Till.    lil((l>IIVI.   lil.WPS 


m 


ihr  |it'i!innl(ilit>  ol  llic  iiiiHiii  Imiilct  III'  lln  111!  Tills  cliaiige  in  iiirnHn- 
I'llitv  niny  lie  <lf|H'iiilciit  upnn  iiltcratin.is  in  surfncc  tension  lirought 
iilioiit  by  flu'  niinrntidii  uf  cIciMml^  lis  I<i  •}■•  I  ii-l«  r.  Tlinf  sucli  a  iiilKra- 
lioii  (»f  elcclrnlyli's  dnts  actually  wxv  lifcn  ilciiKinstialfil  Ity  A,  I! 

\hicalliiMi"   wliit  i|o\ilii|)t'i|  a  iiii<T(«-l;  tcHt    fii;    iiiitiLssiniii.   .mil   was 

.ililt"  to  slio"  that  this  rlcclinlytc  ni'ciiii.:.  s  at  tin-  lumen  ImiiiIi'i-  of  the 
irlj  (iurint:  sccri'l m-y  aclivit\,  I'ntassiiim  may  lie  taken  as  a  piototypc 
i.r  rle('tml.\  tes  in  j.'<Miial.  Sii|i|iiiit  is  (livcn  to  this  vi.w  hy  the  faet 
tliat  potassium  always  aei-umulates  at  the  luiihi  "f  the  (•(■i|  throuu'li 
MJiii-li  the  scrrilinii  l;iKis  placi'  In  the  epitlirlium  of  the  small  intes- 
tine, t'oi-  instanc-e,  uheic  the  euiienl  ({oes  in  the  opposite  iliicction  to  that 
III  u'liincl  eells,  the  ai'i'uiiiulilt ion  ot  piitas^iuni  oeenis  at  Ihf  puition  of 
the  ei'li  111  \l  the  liasemiiil  menilntnic 

Aiioliiei-  piissi|iilil>  is  thiit,  uheii  the  lilainj  lieeniues  mole  aetive,  (he 
iiioleeiiles  present  ill  the  eell  lieeome  lilokeil  down  into  siiiMilei  moleeiiles 
.Hill  so  raise  the  osmotic  jiiessnic  of  the  cell  content,  with  the  result  that 

■  ,itef  is  altiaeleii   from   the   hlood. 
When   the  (iland   is  excited   so  that    the   :i/;ioif7f »  iiidiiiili  x,  as   well   as 

viiter  and  salts,  are  secreted,  the  priniarx  chaiii:e  appears  to  involve  the 
'-•liiiiiiles  only.  Those  near  the  lumen  swell  up  hy  nlisorliiii(r  water,  and 
hecome  converted  into  spheres  in  wliicli  sjilts  are  dissolved  in  smaller 
I'loportioiis  than  exist  in  the  lymph  hathins;  tlie  eells.  These  swollen 
structures  are  then  rupliiied  al  the  periphery  of  the  cell  and  dischar<ied 
inio  Die  lumen.  This  disehart;e  of  a  lliiid  containinj,'  fewer  saline  eon- 
•■titiients  tlian  the  eell  or  siirrouiidiii!;  hlood  plasinn  hrinsrs  ahout  in- 
rr.jis.'d  eonceiitriitioii  in  the  remainini:  parts  of  tlie  i-ell.  a  iirocess  whieli 
l"iNsilily  is  assisted  liy  a  lireakiuif  up  of  molecules  in  the  protoplasm  itself, 
■iiid    which   e.iuses   an    increase   in   osmotic   presMire    with    a    ciinsci|ucnl 

!lo\v  of  wjiter  from  the  l,\mp]i  to  th lis  and  therefore  from  'he  hlood 

I'l  the  lymph. 


OTHER  CHANGES  DURINQ  ACTIVITY 

\Vhnte\ir  ina\  he  the  nature  of  the  |ih\  siolouical  chanues  (li.it 
lie  respoiisihie  for  the  secretory  activity  of  the  c-ll,  the  fact  stands  out 
;irominentl\  that  a  eoiisideralde  expenditure  of  enei't.'.\  is  entailed.  This 
>  iniiiea(ed  hy  (lie  fae(  that  considerahly  Itiisier  i|uan(itics  of  oxyueii 
fe  (aken  up  hy  the  siliiiid  when  i(  is  in  an  active  state  than  when  jit 
"'■St.  Thus,  the  iiriiij' <i  I  ii)i>nn>i)itiiin  of  the  restiie^  siihmiixilhiry  •rlaiid 
"I  thf  eat  may  lie  increased  five  times  dnrin<;  aetive  secretion  On 
account  of  tliis  increased  oxytreii  eonsiimption  it  is  not  surprisiim  that 
it  should  he  found  tliat  the  seeretorx  activity  of  the  cell  is  irreatly  im 
■laiii'd  tiy  a  deficienev  in  owsren. 


m 


41 


i>i(ii>rinv 


TlicM'  iiitivc  pniccsscs  iiiriii^;   in   the   ul.nul   ^^  |„.|i   it    is  rxcitnl   t.. 

MTirtf    iiiv    ;i>MH-iatr.'     with    .•luiim.N    in    clrctiir    if!i<-li..il    alui    in    llir 

^"''"' '■  ""'   «l-""l-     Til"  <l<rlrii-  rl„ni,/,s  Imv.-  Ii.    ,i   ii.ost   .'xfrnsivrlv 

^tll.li.,|   ii,  ....ih.viioii   uith   111,,  siiliviii-y   irhiM.I.     CiuniMi   iiinl   Catl.-I."  I,\ 

••..iiiK-i-liiiif  a   uiilv! iiK'ti'i'   with   ii.iMixiIariy.il.l.'  ,-1..,m     Avs,   mu-   ]A»vr.\ 

•  '11  the  irlaiKl  anil  the  otliir  on  M.i(;lil...iin>{  roMncMivc  (issue,  ammv  nlilr 
tn  slinu  that  uith  .arh  pciio.l  ..f  .irtivf  s.'crHi.Mi  a  .•niirnt  of  action  uas 
s<t    n|i.     This  «as  liist   .Mscomt,.,!   ]n    jfos,.    h:  a.lfonl   .m.l   Mavlisx,   ami 

''""'   ' "   <'f"''l'iillv   stuclini    l.v    (;,.srll."     That    Ih.-   ..l.Hii,.   ..nm-'nt    is 

.l.'lini(..|,\  assorialr.l  uith  thr  s..crrtion  of  saliva  nmi  is  imt  ransc-.l  l.v 
Ih.'   \as,-nlar  rhanu'i's  whi.-h   nsually  aiTcmipanv   this  art    ,.  ,is  shown   l.y 

i'"'    Mir.'nr.-    « li.i,    th.-    I.loo.l    supply    was    shut    off    tVoin    tii.>    irlan.l, 

MM. I  l.\    its  al.s.ii,',.  x\h..M  then'  was  no  sccicti. vrii  lli..u-_'h  the  vascular 

.•haH._'..s   \\v\v   l.n.u'.'hf    al.out  :   neither   is   tli.'  eleetrii-  .hanne  ijue   to  th.^ 

"■ "'   "'■  ""''I  iil<"ii:  Ih.-  iln.'t.  as  .•vitl.-n.'.^.l  l.y  ii     p.-rsiMei afl.r 

ligation  ..I"  the  iliiet. 

With  rtiranl  to  ehani;.'  In  mhim, ,  it  niijrht  he  e\| te.l,  ..ti  ) oimt  «i 

Ih.'   irr.ater   \  as.-iilarity   ..I"  th.'   -rlan.l   i •inpanvin':  a.-livily,    that    this 

"""'"'   '"••"■■•i-<'.     "M    II .nirary.   h.-u.'v.'i.   it    has   h.'.'n   shown   f..  .!.• 

.•r.'as.',  Ii.'.'aiis.'  ..I"  the  larL'e  ipiantity  of  llui.i  sc-r.-t,.,!  I'r the  -rlan.l  .'ells 

''"'i''  ''I'-ti r  l«..  .//•/«/>  on   III.'  L'lan.l  .'ells  is  ..I"  .-onsi.lerahle  physi.i 

lotrie  ii.,p,,rlaM,-,':  that  ..f  atro|.in.',  « hi.-h  |.araly/.'s  Ih,.  s,'er,'li,.u.  an-l 
that  ..!'  iiihi.Mrpin.',  uhi.'li  stimnlat.s  it.  \V,'  shall  se,.  later  In.w  this 
iMr..rni,iti..ii  may  I,.'  usi.l  in  \\..rkini.'  ..ut  the  e\aet  me.-hanisni  ..f  fli.' 
.liff.'ri'ot   "_'laii.ls, 

lrn|...it,inl   .-l.s.rv  alioiis  .• rnin-r  /A.    r,  ^(^"-.«^/,,■/l  „f  ,,l,iii,liil„r  arlir 

ilif  1,1  Ih,  1,1, „„l  siii,,,l,i  hav.'  I>,',.||  m.l. I.'  l.y  exi.,'rini,nts  in  whieh  jrlan.K 
n.'r.'  artilleially  p.'rfiis.'.l  ..ntsi.le  the  l.,,.ly.  Wh.'u  the  sul,maxillar> 
L'lan.l  .if  III.'  .!....'  is  perfus.'.!  with  ..Nyu'enal.'.l  K'in-.r's  solnti..n,  stinmla 
ti.in  ..I"  its  n.Tv.'  supi.ly  .l..es  M.>t  pii..|ii.-e  th.'  usual  si.'reti.in,  liul  it"  tli. 
|{iM-er'ss.,luli..n  is  MiiN.'.i  uith  l.ln.Hl  plasma.  Ih.'  n.'rv,' stiniulafi.in  has  it. 
nsual  .'R-.'-t  for  a  sh.,rt  tin..-.  Allhou-h  n..  s.-.-n'tion  -nvurs  wIi.'m 
oxyu.'iiat.'.I    |{iM..'..r\    so|uti.>n     is    [.erl'iise.!    alone,    the    usual    vas.-ulai 

ehaiiL'.'s  still  -iir  in   th.'  u'lan.I.     Th.'   nsuils  se.'ni  to  iiuli.-ale  that    th.' 

I"'''^''"' r  '*'""<■   e..nstitui'nl    of  till'   l>ioo.|    plasma    is  essential    f..r   th.- 

.'hail-.'  ill  the  (.ermeahilitv  ol"  tl,.-  eell  w.-ill  n.'.'ess.-iiy  for  th.'  pr...'ess  of 
si'.-r.'ti.in  Similar  resnlis  |ia\.'  h.'.'u  ..htain.'.l  ,lnriii-r  artificial  perfiisi.ni 
.if  the  paii.'i'cas  wh.'h  secrelin   vvas  lis., I  ;is  th.-  slliiiulns. 


CONTROL  OP  GLANDULAR  ACTIVITY 

IlaviiiL'  ..iiiliti.'.l  the  ir,.,icr;,|  nature  of  the  ehan^'es  ..c.-tirrin-;  in  jrlii'i'I 
•.■lis  durin;,'  their  a.-tivity.  we  niay  n.iw  pniceeil  to  stuiiy  the  iialuie  of 


i 


Center  for  ^       , 
traniil  secretory  fibers 


rV-^.  ^ 


Fdcial  nerve  mvUL 

Cerebellurf,^:.^^^ 
Cloisoptiarvnqeal  nerve  -.-  j  - 

Medulla  oblonqsta^^-*-' 
Parotid  gland 


Center  for        ^  ^ 
vasodilatorner'ves- 

,'     .    ^OfiC  ganglion 
>;'l?^r'l^'^''t  /^finilunar[Qaiierian)qanqlion 

'     (Nv)  ///t''o''rfa  fympani  nerve^, , 

^^"J/A-imm  superficial  '^ 

-'      ^   petrosal  nerve    -■^Ptrotid  duit 

^    ^Inf  intM  div  NV  (Jf»ojonj) 


Yen 


r, 

nerve        I  , 


Sublingual 
■J 


M'   /     Electrodes^ 
'    /{imtll  amount  of  ihick  sali 
./       vato- constriction  ) 

Vaso  constrictor  fibers 
Sympathetic  secretory  fibers 


duct 

,  Bartholin's ) 
Lingual  nerve 
Chordo-lingual 

triangle 


^       Electrodes 
{Large amount  o/  thm 
,  1     salila. 

'iubmatillarv  vaso  dilatation 

glana 

Sublingual  gland 


•Outgoing  sympathetic 
rami  communicantes 


Post  ganglionic  fibers  are 
dotted  thus  — 


•I,    ,l..u 


MICTOCOPY    RESOIUTION    TiST   CHART 

ANSI  ond  ISO  TESt  CHART  No    2 


32 


12,5 
2.2 


t  ^  m 

1.8 


1.6 


^     APPLIED  INA^IGE     Inc 


...     <r■,^^'      Srw     ■■■N 


^mPTj 

il  M' 

1f  !  1 

rt 

1 

1| 

if      '" 
1 

i 

i'^if^L 


'llYSI(ir.(KiY   OK   Till-;    DKiKSTIVl-;   (il.ANKS 


423 


1h(>  process  l>y  uhicli  tliis  };laiitlular  activity  is  cuiitnillcd.  Two  nu'chau- 
isnis  (it  control  arc  known:  (1)  liy  tlic  ncivons  system,  anil  ("J)  liy  means 
of  liormones. 

Nervous  Control.— Control  tlin>u<jh  the  ner\oiis  system  is  most  marl<e<l 

indeed  it  may  })p  the  only  means  ot"  conti'ol — in  jrlands  uhidi  l>;ive  to 
])i-oducc  tlieif  secT'ction  i)ronii>tly,  wlicfeas  lioiinone  control  i)re- 
dominates  in  those  in  which  ])roni))t  i'han<res  in  se<'retoi'y  activity  ai'c  not 
re(|iiirpd.  Thns,  nervous  control  alone  is  (iresent  in  tlio  salivary  jtlands, 
whereas  liormono  control  is  |>redominant  in  the  |)ancreas,  intestinal 
fjlands  and  liver.  The  gastric  !,'iaiids  are  i)artly  under  nei'vous  conti'ol, 
and  i)art]y  under  liormone  control.  It  should  he  pointed  out  here  that 
the  KJands  of  tlie  body  oilier  tlian  the  digestive  glands  arc  also  suh.ject  to 
nervous  or  liormone  conti'ol  according  to  tlie  jiromittncss  with  which  they 
ai'e  re(piii-e(i  to  secrete.  The  lai-hi'ymal  and  sweat  glands,  and  the  venom 
glands  of  reptiles,  for  exami)le,  ai'c  i)?actically  entirely  under  ner\ous 
control,  wJiereas  most  of  the  ductless  glands,  with  the  excei)tioM  of  the 
adrenals,  are  mainly  under  the  influence  of  hormones. 

The  exact  nature  of  the  nervous  control  of  glandular  function  has, 
therefore,  lieen  most  extensively  stiidied  in  the  salivary  glands,  and  that 
of  the  hornionic  in  the  jiancrcas.  With  regard  to  the  salivarx'  glands, 
the  following  jxiints  are  of  imporfance:  Their  nerve  supply  comes  from 
two  scMirces:  the  cerehral  autonomic,  and  the  sympathetic  autonomic 
(see  page  ST7).  These  two  ncr\i'  supplies  have  usuallx  an  (i|)positc  iiitli! 
ence  on  the  secretory  activity  of  the  glanils,  and  very  f,'ei|uently  also  oil 
the  vascular  changes  that  accom])any  secretory  jictivity. 

On  account  of  its  ready  accessihility,  the  sulnnaxillary  gland  in  tlie 
dog  and  eat  has  lieeii  most  thorouglil\  investigated.  The  cereliral  auto- 
nomic nerve  in  this  case  is  re])resented  hy  the  chorda  tympani,  and  the 
symiiatheti<*  autonomic  by  postganglionic  fiheis  that  run  from  the 
siipi  'ior  cervical  ganglion  to  the  gland  along  its  Mood  vessels  (Fig.  14.")t. 
.\tlcr  tying  a  cannula  into  the  duct  of  the  gland,  it  \\\\\  lie  found  in  IIk 
iloij  that  stimulation  of  the  chorda  t\iiipani  ))rodnces  an  inimediate  and 
ahuiidant  secretion  of  thin  watery  saliva  accompanied  hy  a  marked 
dilatation  of  the  Mood  vessels  of  the  gland. 

That  this  secretion  is  not  depeinlent  on  the  vasodilatation  is  easily 
shown  liy  repealing  the  exiteriment  after  administering  a  sufiicient  dose 
of  atroj)iiiP  to  paralyze  the  secreting  cells.  Stimulation  of  the  nerve  then 
l)roduces  a  vasodilatation  hut  no  secretion.  The  same  conclusion  is 
arrived  at  hy  an  experiment  of  an  entirely  different  nature;  namely.  Ity 
ohserviiig  the  pressure  produced  in  the  duct  A\heii  th<'  chorda  tympani  is 
siimulaled.  This  Jiressurc  rises  ciirisitieraMv  ahove  thai  in  the  arti'ries, 
SO  that  no  such  physical  jirocess  as  mere  ditTusioii  can  he  held  accountable 


M 


M' 


r  ■  1  : 

■ ,.  11 


i. 


424 

for  tlif 


i)i(ii;.vrn)\ 


mil. 


•  liiiwii   from  tlioso  results 


If  s.M.rofion,  nn.l  tlinvforc  vaso.lilatation  alone  enii  not  be  rospon 
sible  for  It.  It  the  sympatlietie  nerve  supply  is  stimulate,!,  a  verv  seantv 
tliiek  seerelioii  tak.'s  plaoc  aeeompanied  i)y  \asoennstrietioii. 

Repetition   of   tliese   experiments    //,    //,,•   rat   yields  ditTerent    results 

partieularly  with   lv.^■„•d  to  the  iulluen. f  the  sympathetie,  a  eopi„n< 

seeret.on  he.nu'  |.rodiu-..,|  i.y  sueh  stimulation.  Th,.  histolo.'ie  eiiaiM'rs 
p.odtuvd  in  th..  -daud  eells  are  marked  after  sympathetic  stimulati-u, 
iHit^  very  shfrht,  if  present  at  all,  after  ehorda  stimulatir 

The  outstandinir  i'( lusinii   hImcIi   mav- 
is that  tw.,  kinds  of  seerel„ry  aetivity  are  me.Iiated  throu-h  the  nervs 

'" •■'"'"'"-  ■■'  "''"   "'"'■'•>•  seereliun,  eontaiuin-  onlv  a  small  pereenta- 

ol  o,M,an„.  ,„att,.r.  and  the  other,  a  thiek  visei.l  seeretiou   with  a  lar-e 
«mou.,t  of  ..r.ranie  material.     To  explain  these  dilTerenees  the  hvputh'- 

MS    has    heen    adva. 1    that    there    an.    really    two    kinds    of    seVre.,,,. 

lil'.'is.  ealle,!  snr.lor,,  and  In.,,!.;,:  the  f.un.er  havin;;  to  ,1,.  with  llii. 
seer,.|,„M  of  water  and  inor-anie  salts,  an,)  the  latt.'r  with  the  seer..tioM 
ot  or,Mn.e  matter:  i.e..  with  the  extnisio,.  of  the  /vmofreu  frranules 
<--''nu  authors  ,[.anudey)  l,elieve  that  sueh  an  hvpothesis  is  unneees' 
,'•'•  "'"'  "'•■"  ""■  •'"1"">''"<  ''snlts  are  ,lep,.n.|ent  up.m  the  eoueomitani 
Hiaimes  ni  the  hl,„.d  supply  produeed  I.y  ,sf imulatiuir  one  or  ..ther  uerv,- 
Tli.-.t  there  are  really  dilVerent  kinds  of  true  seeretorv  fihers  i.  lu.wevr 
'•vi.h.nt    Iron,    the    follnwin-   .■xperiment.      If   the   duet    of   the    .'lau.l    is 

;;""'.■'", "  ,""•"•"-'•'   "  "^'"'"   i"   ^'"^  -I"vk.  seeretiou  of  saliva  "throud, 

""'  '''•"'"  '■""  '"'  ""' '  h   Phwiu-  various  suhstauees  in  the  mouth   sn,!: 

as  meat  powder  or  hydroehlori,,  aeid.     If  the  ..xperimeut  is  performed  n, 

:'"•  '  "  "••'•^    "'•"  ""■  '•! '"""   ""-"nsrh  the  udand  eau  I hserved    it  will 

'"'  '^"""''  "'"^  ""■  ^"''■^•'  I"-""'' '1  •■y  theslimulati.u.  with  the  meat'powde, 

'-"■t'ims  a  very  mue],  I,i,d,er  pereenta^^^  of  orjrauie  material  than  does  tha- 

'','■•"'" '   "'"•"    •'^''' •''"'•!'•   ="'i'l    is   tl,e  stimulant,   whereas  the   vaseul.r 

Hianjr.s  u,  the  -lan.l  an.l  ihe  iuortrani, nstituents  of  the  saliv,  -ire  tie 

same  n,  hmh  e.ises.  Sinee  stimulation  of  the  ehor.la  tvmpaui  eauses  tie 
seeretiou  of  a  wat.My  saliva,  while  that  eaused  I.y  stimulation  of  th. 
s.vmpathetie  is  thiek,  it  u.id.t  he  thouud.t  that  tho  .seeretorv  fibers  we>, 
-outanied  n,  tl„.  former  an.l  th..  tn.phie  fibers  in  the  latter  nerve-  th-,r 
this  ,s  not  tl„.  ,.as,.  eau  1...  sho,-,  n  I.y  a  repetiti.,,,  of  the  ab..ve  experim.n. 
'•;  annuals  fr.uu  whiel,  th..  snperh.r  eervi.'al  iraufrli.,,,  has  b.-en  rem..v..,| 

"."'  """"■  '■'■""""  "'■' '■■'i'"''l.  iN.li.-atin-  that  the  eh..r.la  tvmpaui  .-on 

taiiis  both  s...-r..tory  an.l  tro|.hii.  libers. 


rifAI'TKR  XI. IX 


IMIYSroLOdY  OF  TIIK  DKiKSTIVK  (ILANDS   ((.nitM) 


THE  HORMONE  CONTROL 


This  is  ('xliil)itc(l  jifst  in  tli 


le  case  of  tlie  paiicri'iis. 


The 


lent  tlciiionstratiiis;  that  this  jrland   is  not  primarily  dopcnilfiit    i 


criu'ial  I'xpcri- 


ipoii 


nervous  impulses  for  the  control  of  its  aetivity  was  performed  hy  Hay- 
liss  and  Starliiifr.-  Startinsr  with  the  well-known  fact  that  the  applieation 
of  weak  aeid  to  the  duoflenal  nmeons  meiiihrano  excites  secretion  of  \y,\u- 
creatic  juice,  these  woi'kers  arcfully  se\ered  all  the  nerve  connections  of 
a  |)ortioM  of  the  duodenum,  and  found  on  ajrain  applyinfr  aeid  to  the  mucous 
iiienilirane  that  tl:e  secretion  persisted.  To  explain  this  result  the,\  postu- 
l.-ited  that  the  acid  must  cause  some  suhstance  to  he  liherated  into  the 
lilood  stream,  whicii  carries  it  to  the  ;.aMcieas.  the  cePs  of  which  it  then 
excites  to  activity.  To  test  this  hypothesis  they  scraped  olT  tlic  mucous 
niciiihrane  of  the  duodenum  and  ground  it  in  a  mortar  with  weak  hydro- 
chloric acid  (n.(i  per  cent),  and,  .-ifter  lioilinir  the  solution  so  as  to  remove 
the  i)rotein  ;ind  ncjirly  neutrali/injjr  it.  they  obtained  a  llui<l  which,  when 
in.iectcd  intravenously,  immediately  caused  ji  copious  seci-etion  of  pan- 
i-rcatic  Juice. 

Acccmipanyin-r  the  secretion,  however,  a  marked  fall  in  arterial  Mood 
liressure  was  ol)sei-vcd,  makin;.'  it  possihl,.  that  the  secretion  mi>.'ht  luive 
i>cen  line  to  a  vasodilatation  occurrinjr  in  the  pancreatic  hlood  vessels.  To 
eliminate  this  ])ossil)ility  they  pre|>ared  .in  extract  that  was  free  of  the 
dc|)ressor  suhstanees  liy  exlrai'tinjr  inti'stinal  e|)itliclinm  without  any  of  the 
suhiiiui'ous  tis.sue.  The  resiiltiiiir  exti'act  had  merely  the  secretory  efTect 
and  produced  no  fall  in  Mood  jiressure.  This  secretasioirnary  sulistance 
tliey  named  srrrrlin. 

Further  evidence  that  the  action  i  ''  secretin  is  independent  of  the 
depressor  sulisfances  has  Iieen  olitained  l>y  takiiiu'  ailvaiitajre  of  the  fact 
th;it  the  depressor  sulistance  is  more  soluMe  in  aleotiol  than  the  secretin. 
II  an  acid  decoction  of  duoileiial  mucous  memhrane  is  jioured  into  ahso- 
liite  alcohol,  a  prceiiiitate  is  formed.  If  this  pre.-ipitate  is  redissolved 
in  water  and  rei)recipitated  several  times  liy  alisolute  alcohol,  then  after 
drying  a  white  jiowdcr  is  olitained,  which  is  easily  sohdile  in  wati-r.  The 
fesultiiifi  Solution  iu.n'Cteil  iiitraxenously  lias  a  powerful  secretorv  action, 
Init  produces  no  efTect   on  Mood  pressure.     The  concentrated   alcoholic 


:i:  ; 


^    'il!^! 


;;i 


4::(j 


imihiSTION 


il  ,.  ! 


4 


1 

;    1- 

!   1 

1 

i-  i 

'^ 

ff.ir  I' 


^!in  1 


liJ 


'""""••  •;•'  ""■  ""'«■'•  l-'-".l.  «h,.„  si„.ilM.ly  i„jc.H,.,l  „nMl„,.c.s  a  ,n,.,k..,l  fall 
"I  Moo,|  pirssuiv.  h  is  lK.|i..v,..l  that  this  ..fT..H  is  ,liu-  to  Ih,.  avtio,,  of 
/i-i.M..la...lvI,.thvIan.i.u>.     A  very  stron;.'  preparation  of  MM-rHi,,  ..,»  also 

'".'"■"'""■"''  '•>•  ""'  ""••""""  "'■  "" Hi  hai.llaw/  ul,i..|.  .l..p..M,ls  on  pr. 

«'i|ntatioii  liy  ini'rciiric  diloridc. 

S,...rHiM  .Iocs  no.  ,.xis,  ,,r..fornu.,l  i„  tlu-  rpithclial  ,.,.|ls,  as  is  shown  hv 
tUv  tart  that  an  .xirart,  nia.li'  with  neutral  saline  solution,  does  not  as  a 
"•III.',  hav  any  s,..-r..t..ry  a.-ti.m  uh.Mi  in.j.M.f.Ml  intnivnouslv.  S,.n,..titn.s 
a  sl.sjht  seereti.-n  may  he  pr...ln.-e.l,  hut  this  is  prol.al.lv  t..  he  explain..! 
I.v  the  laet  that  s.,n.e  se.Tetin  remains  I.ehin.l  in  the  cells  as  a  result  ..f  a 
I.re.....ln,u'  phase  ..I'  a.-tivity.  If,  on  the  ..ther  hand,  the  al.ove  neutral  or 
sli-htly  alkalnie  opal..s.ent  s..luti..n  of  the  mu.-..us  meml.rane  is  l..>il..l 
uith  a.-i.I,  M.eretin  may  l„.....nu-  .level..p...l  in  it.  The  interpretath.n  put 
'iP'Mi   Ih.'se  r..sults  is  that  a  suL.stanee,  ealle.l  prosa-ntln,  e.xists  in   th.^ 

.•pitli.'hal  .-ells,  an.l  that  this  I., mes  eouverle.l  into  secret  in  l,v  the  acth.n 

;•'  "-'"I  ">'  Hu.  ..ells.     The  secretin  thus  pro.lu..,.,!  is  then  taken  up  I,v  th.. 

'"",  •  "",',"'  "'■  '*  '''"^^'''^  "'<"'  ""'  i-it.'.stinal  , al.  I.e.-au.se  the  fr...  l,..'r.lers 

"I  the  cells  are  impervh.us  t..  s.-.-relin.    That  this  is  actual. v  the  .-ase  has 
'•".M,  sh..wn  l.y  (in.Iinj;  ,i„„  the  intr<.,lu..ti.,n  of  neutrali/,..|' .secretin  s..lu 
th.n  nito  the  .luo.icnum.  ..r  ..Ihcr  parts  ..f  the  .small   inlestim.,  ,l..cs  n.,1 
cans.'  a  .s.-cctLm  .>f  pan.-r.'atic  juice. 

We  l<n..w  pra.-ti.-ally  n..thinfj  e..nc,.rnin-  the  ./,.  miral  mtlmr  of  sarrtin 
I.nnsr.s..lul.le  in  al.oul  !M»  per  .-cut  al.M.hol  an.l  in  fairlv  weak  aci.is  it  .-an' 
not  helonsr  t..  any  of  the  Letter  ku..wn  sroup,  of 'proteins.  As  it  is 
roa.lily  .lilTusil.lc  thr.msrh  par.-lnncnt  meml.rane,  it  can  n..t  he  of  verv 
e..mpl,.x  siructur..,  an.l  as  it  uith.stan.ls  heat,  it  cnn  not  he  an  en/.vm." 
It  rapi.lly  .l<'t..rM. rates  in  strength  in  the  presence  ..f  alkalies. 

Auy  aci.l  wh.M.  applic.l  to  Ihe  mu.-..us  meml.rane  is  ..apahle  of  pr..<lncin.r 
scMTtn.,  an.l  s,.  are  certain  ..ther  sul.slan.-es.  su.'h  as  mustanl  oil  Waterv 
s..Iuti..ns  ..t  .sacchar...se  ..r  urea,  when  rul.l.ed  up  with  the  .Iu...lenal  m.n-.sa 
in  a  nu.rtar,  i.r...luce  secr..tin  s.,luti..ns  of  varyin-  activity,  I.ut  th.>y  .lo 
U'.t  in  the  l.vn.^r  animal  excite  pancreatic  .secreti..n  whe..  applie.l  U>  the 
•liKHl'Mium.  Se.MTtin  is  very  susccptil.le  t.,  .l.-structi..n  I.v  such  .lijrrstivc 
onzymes  as  th.,se  present  in  the  i.an,.reatic.  jrastric,  an.l  intestinal  jui.-cs 

Imt  .s..cretn,  ,s  present  in  the  1.1 1  when  aci.l  is  in  c..nta..t  wiih  th.' 

"""'''""'  "" '*"  '•'"*  '•<'*'"  '^•'""'1  I'.v  tl.o  fact  that  inject i..n  into  a  n..rmal 

<!(.-  ..f  1.1.,.„1   fr.,m  one  i.i  which  secretin   f..rmali..n   is  Koing  on   (as  a 
result  of  ac.,1  in  the  .lu...l..num),  excit.'s  pa.u-reatic  .secretion. 

Th.-  pancn.ati.-  juice  pr..,lu.-.>d  l.y  the  injecti..,,  of  secretin,  like  that 
uhu-h  IS  pr...lncc.l  un.lcr  normal  e..ndili..ns.  .l..cs  n..t  cntain  any  activo 
trypsni,  hut  nistea.l  .-..ntains  its  j.recursor,  trypsino-en.  This  i.ecomrs 
converte,!  n.to  trypsin  in  the  intestine,  I.eing  activate,!  by  contact  with 


■iivsi()i,(KiY  (IF  Tin;  i>i(ii;sTivr:  (ii.anks 


427 


I  iitcrokiiinsc,  <'iii  on/viiic  pirscnt  in  tlic  iiitt'stiniil  Juice.  My  siicli  a  iiicclian- 
isiii  the  niUfosa  of  tlic  paiici'oatic  diK-t  is  pnilcctcd  a<;aiiist  autoiligcstio'n 
liy  trypsin. 


NERVOUS  CONTROL  OF  PANCREAS 

I'ricir  f(i  tlip  (liscovci'v  of  .scciclin,  Pavlov'  anil  iiis  pupils  liad  iinblislicd 
iiiimorovi.s  oxporinu'iils  piirpoi'tin^  to  sliow  tiiat  tlii'  scci-ftioii  of  paiici't'atic 


I'lK-  14'..  I'iirurt.ttit-  .liini  >t.iiiU'l  with  liriu.it. ■wliii.  'I'lu  .uim  ;it  tin  t..].  .iii.I  t..  the  )<  tt 
.1  ttu-  liKuri-  ate  fri'in  :i  if>Iiin;  Kl.'tf'l.  Il.">f  to  llii-  iiKhi  lifinK  tr..iti  (.iic  tliat  It.i.i  tutii  sriittiii^; 
l..r  <t\tT  thric  Imurs  as  a  rt  suit  ot  aci.l  in  tin  lin.t'it  itiim.  Tin-  l'.wtrni>'>t  lik'iir*  is  ffmi  a  K'.i'i'i 
'■'■'  vaKus  lurvi-  Mi|.|'ly  ..I  ulu.li  li.i.l  Iphii  slnniiKilnl  nil  aii.l  ..n  l,.r  smi.ii  li..iits.  N.ili  that 
111.  /yiiioKfii  Kfaiiillts  art-  r\ttM.h..!  ..illy  afttr  ya^'.is  aitnity  l.iil  n..l  aftt  r  siirrtiii  aitivMy.  iImimii 
I'. i'. kill.  Kiihasi-hkin  airl   SsawitM-h.l 

.jiiioo  is  Odiit rolled  tliroiij;li  the  vasr\is  nerve.  Tlie  aiimnnt  of  secretion 
iiriiiliiced  l»y  nervous  stiiiiiilat ion  \\;\s,  however,  nevi'r  found  to  lie  so  lars^e 
us  that  produced  by  secretin,  and  for  several  years  after  the  discovery  of 


iM 


i   HI 


1  >^ 

i  i   ] 

; 

ii\ 

;  1  \  1 

v 


4-JH 


li.  • 


111 


iSi 


m 


It 


i    :.i 


1    ! 


nmisrids 


the  Intt.r  honnono,  m.u-h  d-ul.t  oxi.f.a  a.s  to  f ho  ..orrootness  of  I'nvlov '. 
.  n.m.     As  .„  „,any  uthor  fields  ,.f  physiologi.-  s..ion,..,  investigators  .. 
■"Ptod  ,o  s  ou  ,l,a,  o„o  o,.  ,1,0  ..,l„...  ...eohnniM,.  oi.tainod.  and  thov  .,.,.. 

H  ,  ,„,.|„„.d  ,.,,.. .nsMlor  ,1,0  possihilijy  ,l,a,  i.ofh  n.oohanisms  might  oxiv 

ork    ,n  M  „  has  Loom  fo.n.d  ,hat  if  proper  prooautions  are  ,ak..„ 

•P.'.'.fd  s„„„,la,>,.M  of  ,1,0  va,M.s  none  does  oall   for,h  a  soere.i..,,  „. 

"••""•'•"■••'""  .1'-"   -hieh,   hosidos   l,ei„g  less  oopio,.s   ,ha,i   that    follow!,,.- 


■^if 


II. 

Tin 


Kra,.h,    l,..,v,.   f„r,n..,l   a„.l    n,.r.    I.c     ,./,',"/    LV,,,, 'f^  \^^                           ','T''   •■'•"  "<    '"    ""-•    ■•'-> 
.r,-ar   ,„    ,h,.   ,.|,.ar   ,.n>,„,,,aM„.      ,  ,^r,„„    1^,'kin      KnKi;:!,,'^;!^^:^;;^;;;--''-   -"1   "■-■  -m, 


soere,,,,  ,„jo..„o„    dirf.Ms  ,V,„.,  i,   i,.  „,,  i,npo,-,ai„   fact  that  it  o.„„ai,. 
'-<  trypMnoooM  l,n,  a..,ivo  ,,ypsi„.     Si„..o  ,l,o  ..o-.nal  pancrea,io   ji.i,-. 

;■;"■■""'  "••^■'" r'''-  "'''  '"■'*  ""'"ti""*'.!  iao,   u„„Id  appear  to  ind'ioa.. 

that  vaK„s  e,.,„n,l  „,  ,1,.,  ,„.,,.,al  soero,io„  oai,  not  l,e  an  important  afTai, 
The  vau'us  s,..,...,,,.,,  ..f  „an..,-,.afi,.  juieo  is.  n,..,.oov,.r.  pa.alvzod  l,v  a,rn 


pine,  whioh  has  no  aoti 


H't,o,,  0,1  the  sooretin  meoh 


ai,ism  (of.  Bayliss). 


'I'lie 


IMIYSKII.OiiY    (ly   Tin:    HKJISTIVi;   (il.VNKS  429 

copious  scciTtioii  of  p.-iiicitatic  juice  iiroduccd  liy  secretin,  on  the 


hand,  and  the  scanty,  tliic 


secretion   pi'odnced   by   vajjus  stiniula- 


tiuii,  on  tile  otlier,  calls  to  mind  similar  dilTercnccs  oliserved  in  the  sccrc- 
tinii  of  saliva  as  the  result  of  chorda-t\  nipani  or  sympathetic  stiiiulatiou. 
it  will  lie  i-eineMd)ered  that  from  these  latter  results  it  was  ci'iicluded 
that  there  must  be  secntory  and  troi)hic  libers  concerned  in  the  control 
iif  the  activities  of  frlan.i  cells.  Interestiiif;  corroboration  of  'his  conclusion 
lias  recently  been  obtained  by  lii.iluloi/ii'  ( .niniiiiutioii  of  tin  paiiirfas  fol- 
lulling  secretin  or  nt<iiis  (ictivUi/.  After  the  repeated  injection  of  seere- 
iiii.  it  is  ditlicult  to  observe  any  siyrns  of  fati<.'ue  in  the  cells;  the  zymoKcn 
uianules  renniin  jtraetically  as  nuniei'ous  as  in  a  resting  •;land,  but  in  the 
riear  proto])lasm  of  the  outer  third  of  the  cell,  it  is  said  that  tine  channels 
Hi'  tluid  can  be  seen.  Through  these  channels  water  is  believed  to  pas.s 
irDia  the  blood  towards  the  Iniiicn  and  in  its  course  to  carry  with  it  some 
i><  the  zymogen  -,'ranules,  without,  however,  ehanirin!,'  them.  Thus,  when 
the  5,'land  cells  are  stained  with  eosin  and  oranije,  after  secretin  activity 
suiMC  of  the  zymogen  firannles  riii  occasionally  be  seen  in  the  lumen  of 
tlie  acini  stained  with  eosin  like  llmse  in  the  cell  itself.  After  vajjns 
stimulation  the  appearances  are  dilTerent ;  not  oidy  are  the  !,'ranules  more 
Ifccly  extruded  from  the  cells,  but  they  uMder<,'o  a  prelimiiiaiy  change; 
they  lose  the  property  of  .stainin;,'  with  eosin  and  bee(mie  stained  with 
the  oransre,  at  the  same  time  increasing  in  size  so  as  to  form  vacuoles. 
Tlioso  vacuoles  may  wander  into  the  ductules,  and  when  they  are  present 
iiiro  they  are  .stained  by  orange  (l-'igs.  14()  and  147)  (Habkin,  etc."»j. 

Why  there  should  be  both  a  nervous  and  a  hormone  control  of  the  pan- 
I  reatic  secretion  is  not  clear.  This  i;land,  unlike  the  "rastric  and  .salivary 
'.'lands,  is  not  called  upon  to  become  active  all  of  a  sudden,  and  it  is  dif- 
ticult  to  see  what  could  serve  as  the  normal  stimulus  operating  through 
the  nervous  pathway.  Taking  it  all  in  all,  it  is  probably  .safe  to  eon- 
'liido  that  the  nervous  mechanism  is  relatively  unimportant,  and  that 
under  normal  conditions  it  seldom  if  ever  is  called  into  operation.  Cor- 
I'lhoration  for  this  view  is  afforded  by  the  fact,  above  mentioned,  that 
the  pancreatic  juice  produced  by  vagus  stimulation  contains  active  tryp- 
■^in,  which  is  not  the  case  with  normal  pancreatic  juice. 


i 


i 


-■:|| 


.ir 


'i 


I 


I. 


■Mi 


Hi 


•IIAI'TKK  \, 

I'llYSlol.udY  oi-  Till-:  IIKIKSTIVK  (ILANDS  (C.ntM) 

I'l.  to  fho  pr..s..Mt  u..  li.nr  !„.,.„  ....n.-.Tii.,!  with  tho  phvsiolcKie  „,.ti\i 
firs  ,.t  ,|.u'.st.v,.  ulmuls  in  jf,.M,.|al.  l.ut  im.w  u,-  ...i.st  slu.lv  ...iHi  of  fh.n, 

soimrMt..Iy  ■„  onl,.,-  f,  fi,„l  out  tl. .Miitions  u.i.l,..-  wl,i,.l,  thov  1m „„ 

sfiniiilat.Ml  to  ,„.tivit.v  in  tl.c  normal  pr ss  of  .li^r.-sfion.     Tho'sociTtion 

«'(  •■n.-h  t'liUHl  has  a  .hfinit.-  r assiirn,.,!  to  if  in  th.-  .•oMipl.-x  an.l  h-n^Mh^ 

proross  of  ,li;r,stion.  It  tak.s  „p  its  work  where  the  pro..p.Iin>r  srrrr 
t.on  l,.|t  ofT:  v.i:.,  th.-  popsin  of  ur„strio  jui.-o  .lijf,.sfH  prof.-in  so  far  .^ 
prot..os,.s  an.l  p.ptono;  th,-  trypsin  of  panrrcati,-  juico  then  attacks  th, 
protroses  an.l  peptone,  an.l  the  resnltins;  |.,wer  .le^ra-lation  pro.lu.'t. 
a.-.'  finally  atta.-ke.l  l.y  the  erepsin  .,f  the  intestinal  jui.T  The  se.Mv 
lions  of  th,.  various  «lan.ls  are,  th..ref..r...  re.p.ire.l  in  a  eertain  .leflnii,. 
..nier  they  an-  eorr..|al..,l:  an.!  we  must  ,h.w  ^ive  some  attenti.,n  t.,  tl,. 
pre..,se  e.,n,lition  upon   whieh  the  a.'tivily  an.l  e.ufelation  .lepen.l. 

THE  NORMAL  CONDITIONS  UNDER  WHICH  THE  GLANDS 
BECOME  STIMULATED  TO  INCREASED  ACTIVITY 


aetivities  of  tin 


•!  f  1     ^        I 


To  make   p..ssil,le  sueh   observations  .m   the   normal   ,-,  „.    , 

srlan.ls.  a  preliminary  operation  has  t..  1...  performed  so  as  to  brii..'  t 
•IiK-f  of  the  j,Han.|  to  the  surfaee  of  th,.  l..,.ly  an.l  permit  ,.f  the  observ,, 
tion  ..  ,ts  s..er,.t..ry  aetivily  affr  the  .-.uimal  has  re,.overe,l  from  tin 
""""•"liale  eir..ets  of  the  operati,.,,.  We  „w,.  to  Pavlov-  the  sur-ie , 
le.-hn.e  by  whieh  these  e..n,liti.,ns  .-an  bo  fuKill,..!.  The  jjeneral  prineipl. 
"*  ;•"■  "'P'-nition.  in  the  ease  of  .^Ha.uls  ,,rovi,l,.,l  with  .lu.-ts,  e.msists  i, 

...ak.ntf  a  eireular  .-ut   thnnisrh  the  mi us  membrane  surroun.lin-  tl,. 

op,>nmK  of  the  ,luet   an.l  then,  after  ,liss,.elin!r  the  ,luet  fr,.e,  stitehin 
the  ...ljc,.s  of  the  eut  t.,  the  skin  w.uin.l.    llealinsr  then  tak,-s  pla.-e  with.,,, 
'  "■  "•'■aiat...n  in  the  ,lu.-t  ..f  any  stri.-ture  ,|ue  t..  th,.  eh-atrieial  tissue     \ft, 
th,.  woun.l  has  h,.al,.,l.  the  seereti.u.  ,-aii  r,.a,lilv  be  e..lleete,l  in  a  m.eiv, 
atta,.h,..l  over  th,.  .lu.-t  fistula,  the  animal  bein-  in  everv  other  wnv  in  .• 
perf,.etly  normal  ,.oM,liti.,n.     In  the  ease  of  «lan,ls  not  provided  with  .. 
duet,  other  m,.tho,ls  must   be  adopte.l  l„  eoileet  the  seerelious      Tlu.s, 
will  be  deseribe,!  elsewhere. 


4;!0 


I'llVMIOI/XIV   OK   THK    I>IClf:.STIVK   «I.A\r>S 


THE  NORMAL  SECRETION  OF  8AUVA 


411 


The  diiPt  (iMfiila  cnii  in  tliiM  ens*'  lie  iiiiuh'  »'itln'r  for  tin-  sulmiaxillaiy 
irlnnd,  n-pn-scnlinu  ii  nnicoiis  k'ihhI.  <>i'  tnr  the  pnroti<l,  rcpresi'iitiiin  n 
serous  Klaml.  I'lulcr  (irdinary  ('(iiiditions  tlicrt'  is  very  little  .secretion 
fr<iiii  either  duet.  When  secretion  occurs,  it  is,  of  course,  caused  l)y 
influences  acting  on  n  nerve  center  or  centers  in  the  ineilulla  ohhnijfata, 
the  exact  location  of  which  for  tiie  ilifTcrent  ({lands  lias  lieen  worked  out 
in  recent  years  Ity  Miller."  The  impulses  acting  on  these  ccnter-s  may  he 
transmitted  alontj  afferent  nerves  coming  from  the  mucous  meiuhraiu'  of 
'he  mouth,  nares,  etc.,  or  hy  imini'ies  which  we  may  call  psychic,  trans- 
niifted  from  the  hiirher  nerve  centers.  The  reflex  secretions  caused  hy 
iinpulses  travelini;  by  the  afferent  nerve  from  the  month,  etc..  have  been 
railed  iniituKlilwiiiil,  and  those  from  the  higher  nerve  i-cnfcrs,  coiiili- 
tUtiuil.  With  retrard  to  the  forner,  there  is  considerable  discrimimition 
in  tiic  type  of  stimidus  that  will  be  elTcctivc.  Thus,  if  the  <hnj  for  most 
of  the  experiments  have  been  performed  on  this  animal  is  fiveii  meat, 
a  secretion  of  tliick,  miii-ous  salivii  will  be  observed  to  occur  (submaxil- 
lary (fland).  On  the  other  hand,  if  the  meat  is  drie<|  and  i>ulveii/.ed, 
the  sci-relion  whicii  it  calls  forth  will  be  very  copious  and  watery  (par- 
otitl  ({land).  There  is,  then,  an  obvious  association  between  the  nature 
of  tlie  secretion  and  the  functiini  it  will  be  called  upon  to  perform  when 
it  becomes  mixed  with  the  food.  The  mucous  secretion  called  forth  by 
meat  will  serve  to  lubricate  the  bolus  of  food  and  thus  facilitate  its 
s\\allowin({,  whereas  the  thin  watery  secretion  prodiu-ed  liy  the  dry 
powder  will   have   the  efTect   Oi    washin!{   the   pow<ler   from   the   mouth. 

It  is  evident  tliat  the  mechanical  conditirui  of  tiie  food  partly  deter- 
mines its  exciting  (piality.  Mechanical  stimulation  of  the  mucosa  alone  is, 
however,  not  an  ade<|uatc  stimulus,  for  if  pclildcs  are  placed  in  the  mouth, 
little  secretion  occurs,  but  if  sand  is  pla(  cd  in  the  mouth,  secretion  iinmcdi- 
Jitcly  becomes  copious.  The  nerve  endin;{s  also  respond  to  cliciiiical  stidiiili. 
Tims,  weak  acid  causes  a  cojiious  secretion,  while  alkali  has  no  efTect; 
disagreeable,  nauseous  substances  also  e.xcite  secretion.  The  above  dif- 
ferences in  the  response  of  the  ({lands  acc(n'din<{  to  the  mechani<'al  condi- 
lion  of  the  food  has  been  observed  onl.v  in  the  case  of  the  parotid  u'land, 
increase  in  the  submaxillary  secretion  bein'{  obtained  only  when  actual 
foodstufTs  are  placed  in  the  nmuth. 

The  invest ij{ations  that  have  been  made  on  the  conditions  of  psyehie 
seeretiiui  of  saliva  are  still  more  interestinj-  and  inipoi'tant.  Their  im- 
I>ortanee  depends  not  so  miich  on  the  information  they  <i\\o  us  eoncern- 
inj{  the  secretion  of  saliva  as  such,  as  on  the  methods  they  furnish  us  for 
inve.«.tif»atin({    the    various    conditions    th;it    afl'cct    the    psychic    jinK-esses 


^1 


I  4 


j'  1 

if  i 


I' ' 


4.R! 


lll<)K>4Tlll\ 


iisMMiiit.d  Willi  ilic  takiiiK  ..f  I'ikhI.  It  is  t'lnm  the  psyrlii.-  rafli.T  lliiiii 
rnmi  tlu'  physidlotfio  stiiii<l|Hiiiit,  tht'N'f.nv,  that  tlu-sc  olisi'ivatimis  ai>' 
of  iiiiportiiticc,  I'ltr  tiny  pciniit  us,  by  oli.jcftivc  niflliiiils,  to  stiiily  mi 
iliiiiili  animals  inolilcms  that  xvimld  otliciwisf  lie  lityonil  our  powers  of 
liivcHtiKatioii      Many  of  the  results,  with  iju'ir  lteariii«  on  flie  fuiietii.iis 

of  tlie  lii(flier  iitrvt iitns,  have  lii'eii  disciisse.!  elseuhere.     ileaiiuhile, 

however,  even  at  the  risk  of  repetition  it  may  not  he  out  of  plaee  to  eite 
a  few  of  the  nnist  iiiterestiujf  exjieriuients. 

If  we  lease  a  hungry  aiiiiiial  with  food  for  which  he  lias  a  ttreat  appe- 
tite, a  eopious  seeretion  of  saliva  iminetliau'ly  oeeiiis.     If  we  ko  on  teas 

iiiK  him  without  ifivlni?  him  food,  nnd  rei)eat  this  pi edure  on  several 

sueeeediiiir  days,  it  will  ho  found  that  tiradnally  he  no  lonu'er  resiMinds 
to  the  feasint;  hy  inerensed  salivation.  Kvidently,  therefore,  the  retlex 
is  eoiidifioned  upon  thp  nnininrs  afterward  reeeiviiiK  the  food. 

The  experiment  may  he  Performed  in  another  way.  If,  for  example, 
we  offer  the  animal  some  food  for  whieh  he  lias  no  appetite,  no  seere- 
tion of  saliva  will  oeeur;  hut,  if  at  the  end  of  the  juoee.ss  wv  ijive  him 
api>eti/inn  fiod,  it  will  lie  found  after  re|)eatiii(;  this  pnieedure  on 
several  sue.essive  days  that  the  presentation  of  the  uiiapi)eti/.in(,'  fooil 
ealls  forth  a  seeretion.  He  has  learned  to  ns.soeintc  the  presentation  of 
unappefizin«  food  witli  the  sul>;ei|uen;  t'latifiention  of  his  appetite.  Tiie 
experiment  ean  even  he  performed  so  that  n  definite  interval  of  time 
elapses  hetwcen  the  ai)plieation  of  the  stimulus  nnd  the  salivation:  if 
the  animal  is  teased  on  sueeessive  days  with  food  for  whieh  he  has  an 
appetite  hut  is  not  (riven  the  food  until  after  ten  or  twenty  minutes, 
presentation  of  this  food  will  eoino  to  ho  followed  by  salivation— not 
immediately,  hut  after  the  exaet  interval  of  time  that  had  been  allowed 
to  intervene  in  'he  training  proees.s.  During  this  interval  there  must  ho 
an  inhibition  of  psyehio  stimuliiti<ui  of  the  .salivary  eeiiters  by  other  nerve 
eenters.  It  is  of  Rrent  interest  that  this  inhibition  may  itself  be  inhib- 
ited by  various  foriiis  of  stimulation  of  the  nervous  system  (see  pajro  SoH). 


I  J 


!  ■•! 


THE  SECRETION  OF  GASTRIC  JUICE 

Methods  of  Investigatioc 

There  beinp  no  eommon  duet,  the  seeretion  of  the  Rastric  glands  is  a 
mueh  more  difficult  problem  to  investigate  than  is  that  of  glands  which, 
like  the  salivary,  are  supplied  with  duets.  One  of  the  most  interesting 
chapters  in  the  history  of  physiology  concerns  the  methods  which  from 
time  to  time  have  been  evolved  for  the  collection  of  this  juice  and  for 
studying  the  digesli\e  pioces.ses  in  llie  stonuich.  Prominent  among  the 
problems  confronting  the  earlier  investigators  was  the  question  whether 


1 


I'llVSlOUNlV    UK    Till:    liKIKMTIVr,   tlI,,\M>S 


431 


iIm'  mniii  liiniMimi  nl  llic  j.tip|iiiifli  in  to  i-nish  i>i-  ti  iliiialr  the  I«mm|  or  1" 
;ii-t  oil  it  fhtiiiicjillv.  Till'  (tiTiit  I'iclifli  NcifliliHt  Ki'iiUiiHl I'  iiinl  a  little 
iiitiT  tilt'  Italian  AI>Ih'  S|mllaii/.aiii  (  l7"-'fl-!7!»!li  altai-knl  this  i.ioIpIciu  Ii> 

iIiixIn  tliiil  iiiitifi|>iitt'  those  of  KihliiNs  and  Kiiihoin.    S|ialliiii/aiii  iilti- 

iiiiiti'ly  ilvviscil  the  im'thod  of  s«  allowiiuf  small  |»'ilcirat<tl  woo.lrii  tiilx's 
i  oiitaiiiinif  foodstiilTs  ainl  covcitiI  hy  sinail  liin  ii  l>aus.  Alter  the  hau's 
uci't'  passed  per  reetuiii,  he  t'oiind  that  eoiiMilerahle  erosion  or  diu'estioii 
of  the  food  had  oeeiirred,  Imt  that  the  woodm  tiihej,,  houever  tliiii- 
uiilled  they  iiiiifht  lie,  were  not  crushed.  In  order  to  sn-iire  samples  of 
till'  uiistrie  juiee  free  from  food,  the  only  method  a\Milalile  to  the  older 
iiivi'stinatiirs  eoiisisted  in  swallowing  spoiiu'es  attiiehed  to  threads,  wliieh 
.liter  lieiiitr  for  some  time  in  the  stomiieh  wi'ic  withdiiiwii  and  si|iiee/,ed 
ilr\  of  juiee. 

The  next  ni'eat  eonlrihiilioii  eaiiie  from  this  eoiiiitrv,  w  lii're,  in  l>«:>:i, 
l>r.  n"aiimont.  while  a  siii'_'eon  in  the  sers  iec  of  llir  Amerienii  troops 
loeat.'d  at  Maekiiiaw.  made  olis.-rvaf ions  on  a  CaiiiidiMii  voyajjeiir  li.\  the 
name  of  Ale\is  St.  Martin,  who  liy  the  jirematiire  diseliarite  of  his  '_'iiii 
li.id  wounded  himself  in  the  stomach,  the  wound  inver  healin','  Imt  leav- 
iii-;  a  peniiaiieiit  trastrie  fistula,  heaiiiiioiit  arraiiifid  to  keep  Aie\is  St. 
.Martin  in  his  service  for  several  years,  diiriii!.'  which  time  he  made 
numerous  ohservations  on  the  process  of  ditrestion  in  the  stomach 
nliscrvations  many  of  which  are  of  >.'rcat  value  even  at  the  present  day. 

Hy  none  of  these  iiielhods,  however,  could  a  sample  oj  pure  u'astric 
Juice  lie  secured  while  the  ditjestive  process  was  actually  in  pro'.'iess. 
To  iiiiike  the  collection  of  such  a  saiiijile  possilile,   llcidcnhaiii  de\  iscd  a 

iiicthoi!   of   isolatint;   portions  of  the  stomach    \\all    as   p lies   opcniii'.' 

throuifh  (istiihe  on  the  alulominal  wall.  The  results  of  llei.lenhain 's 
cxperinieiiN  are,  however  open  to  the  oli.jection  that  the  secretion  in 
the  isolated  pouches  may  not  really  <'orrespoiid  to  that  occiirriiiu'  in  tli" 
main  stoniaeli,  since  the  conneetions  of  the  poiiehcs  with  llic  central 
nervous  system  must  have  lieeii  severed.  In  orilcr  that  these  connec- 
tions iiiijiht  remain  as  nearly  intact  as  possilile,  the  KiissiMii  phy^ioloL'i^t. 
I'avlov,'  devised  an  insieiiioiis  operation  in  which  the  pouch,  or'*iiiinia- 
liire  siomacli."  remains  eoiineeted  with  the  main  stomach  thiiiunh  a  i-oii- 
sideralile  width  of  mueiuis  and  sulimueinis  tissue.  The  essential  nature 
of  this  (iiieration  will  he  evi<lent  from  the  accompMiiyiiif;  diairram. 
'Fin.  148K 

The  most  reecnt  investiirations  have  licen  made  hy  Cannon  ;ind  liy 
Carlson.*  The  former  fed  animals  food  impre«:nale(|  with  Kisiiiulh  ^ul>- 
iiitrate.  and  then  exjiosed  the  animal  to  the  x-rays.  A  shadow  i>-- 
produred  hy  the  food  mass  in  the  stomach,  and  from  the  changes  in  the 
outline  of  this  shadow  facts  have  lieen  eolleeted,  not  only  eoneerniii'?  the 


^'1 


liS, 


m 


i  il 


( 'HP.' 
■llP 


<■■!» 


'il'-' 


:,«! 
n 


Wi  ■ 

1    i 
i'  ■ 

iri; 


» 


i'i<;;:sTi()N 

"""VC'ltlCllts    of    ll„.     vis,.US.    Illlt     ills.. 


''"'"if  <•'«•  rate  of  (lischarjfc  oi 


!•:=:• -:i;;:::;;i:t:;.;;::;::-^ 

r;"'-:"!' ' ■'■■ .:^:::  .li{t  :z:z: 


■i'"l   fiuiii  III,,  niain  sl.ii„.i,  h 
1 Ii  "11   llii-  .iIlIcmiihi,,! 


"■'Ii.       ll'.u|.,v.J  II" '..'ir.iii.  .       .1  ....    l^    ih,     .,|„.„iiii,.    ,,|    III 

i;;::;,;;:: ::::;;:  ;;,:;::;■ '■ "■ >'■.'" ^.i... 

The  Nervous  Element  in  Gastric  Secretion 

Til"  (i.-s(  slinn.lMs  t..  ll„.  .s,.,.r..ti,M>  ..f  uMs,,i,  j,,!,,.  is  m.m.vi.m     •         •    • 

;*:'::::;■■"■:;;  ■:;,::7::;;:':r-"'; ■-*'-';::-.= 


■•fll|"«i"iTO-li  irsvlf  „i.  iv  .       ..       ■ "(■"".■M  ii„i„  „  lisii.i 


•ii|i(.,|  llii-.iiisrli  (1|,>  e.soj)lia- 


liji 


i'iiYsi()i,o(iV  (IP  Tin:  iiKiLsrivK  <;i,am>s 


4:!r) 


.j.al  fistiilii.  Tliis  cxiHTiinciit  is  kmiwii  iiK  Hint  of  "sham  fwdiiii,'." 
Within  a  few  inimitcs  aftn-  civiiiu  Uh,<\  llic  s;astric  .iuicc  was  found  to 
h(>  sfcrcti'il  adivcly,  ami  if  tlic  fcc<linu:  i.roccss  was  kcjit  up,  wl'i.-h  coiiUl 
lie  dnnt'  almost  inilclinitcly  since  llic  animal  never  became  sal  islii'd,  the 
secretion  eontiime.l  to  llow.  Thus,  in  one  instance  i'avlov  succeeded  in 
colloctin!.'  ahout  700  c.c.  of  fiastiic  juice  aflei-  sham  tVediiii;  an  animal 
fdi-  live  or  six  hours  in  the  manner  above  dcserilied. 

After  the  stonmi-h  has  emiitied  itself  of  the  foo.l  taken  with   the  lire- 

\ioiis  meal,  it  is  said  hy  I'avlov  t( iitain  only  a  little  alkaline  mucus. 

The  more  recent  work  of  Carlson,  however,  shows  that  this  i-  not  strictly 
the  case,  there  liein;.'  more  or  lc>s  of  a  coiitiiMions  secretion  of  <rastric  juice 
ill  the  entire  aliseiice  of  food.  The  an  ouiit  varies  fnmi  a  few  e.e.  ui>  to 
(10  f.c.  per  liour,  more  secretion  heiiiL'  p  -oduced  when  it  is  collected  every 
live  or  ten  minutes  than  if  it  is  collecte.l  every  thirty  «r  sixty,  thus 
indicatintr  that,  ordinarily,  some  escapes  throii-xh  the  i)yloriis  into  the 
duodenum.  The  secretion  contains  both  pepsin  and  hydrochloric  acid. 
As  to  the  cause  of  this  continuous  secretion,  little  is  known.  It  may  be 
an  pxamide  of  the  ix'riodic  activities  of  the  <liu'estive  udaiids  ib'sciibed  by 
I'.dldyrelT,  or  it  may  in  i)art  be  <lne  to  a  psychic  stimulation  deiiendent 
upon  the  thought  of  food.  That  the  latter  is  inobably  not  the  cause,  is 
iiKlicated  by  tlie  fact  that,  at  least  in  Carlson's  patient,  the  i)sy(diie  juice 
loiild  not  lie  made  to  flow  short  of  siivinsr  food. 

The  sham  feeding  causes  stimulation  of  the  srastric  secretion  throii'.'h 
iiiiimlses  transmitted  to  the  stomach  aloiiu'  the  vaaus  nerves:  for  it  has 
been  found,  in  animals  in  which  the  vaiiiis  nerve  has  been  cut,  that  the 
sham  feedinir  no  lonsror  induces  a  secretion  i.f  irastric  juice.  The  ques- 
tion therefore  arises  as  to  how  the  nerve  center  is  stimulated.  Three 
possible  causes  may  be  eoiisi(b'red:  l\)  iiieehanical  stimulation  nf  the 
sensory  nerves  of  the  mouth:  CJ'  chemical  stimulation  of  the  nerves; 
':^  the  airreeable  stimulation  of  the  taste  buds  and  olfai-tory  ei'.din>:s 
iiiiicerned  in  the  tastiiii.'  of  food.  In  investiL'at  iiiy;  tliese  possibilities, 
mechanical  stimulation  was  readily  ruled  out  by  showinu'  that  men' 
takiu!.' of  solid  matter  in  the  month  did  imt  excite  any  secretion,  althouuh 
it  misiht  cause  a  llow  of  saliva,  ;^^ere  chemical  stimulation  could  not  be 
the  cause,  for  no  secretion   was  imbieed   by   iilaein^'  substances  such   as 

; tic  acid  or  mustard  oil   in   the  mouth.     I?y  exclusion,  then,   it  would 

ai)pear  that  the  ade(|uate  stimulus  must  consist  in  the  airreeable  stimnla- 
tinii  (if  the  taste  buds,  etc.-  that  is  to  say,  in  the  iinifilirdll'Hi-  of  upprliti . 
Further  justifieatimi  for  this  conclusion  was  leailily  secured  by  notini; 
that  foodstuffs  for  whi(Oi  the  animal  had  no  )iarticular  desire  (u-  appc- 
ite  failed  to  excite  the  secretion.  Most  dou's,  for  example,  althouu'h 
they  may  take  it,  are  not  particularly  fond  of  bread,  and  when  feil  with 


,*.[[ 


'\vn 


i 


m\ 


i)i<ii:sTio\ 


ill 
Hi 


it,  tlics.'  niiiiiiiils  dill  not  j)i„,lu.-..  iiiiy  mipHito  juice.     In  on,-  nniiual  tlijil 

^''""•'''  >si.I(.niMo  likinjj  for  l.rcad.  aftivc  spcirtion  occurred  whc-n  lie 

A\as  (V(l  with  this  t'oodstufT. 

Pavlov  I'urtli.T  noted  (hat  usually  it  was  not  necessary  actually  to 
allow  the  aiiinial  to  take  the  foo.l  into  his  mouth,  l)ut  that  mere  teasin- 
with  savory  foo.l  was  sutllieient  to  cause  the  secretion,  and  that  in 
hi-hly  sensitive  animals  even  the  noises  au<l  other  events  usually  asso- 
ciated with  feediii'-'  time  were  suflicient  to  excite  the  secretion.  In  tlie 
••ase  of  ;,  hun-ry  animal,  the  mere  a|)i)roach  of  the  attendant  with  food. 

'"'   ■'"" "'"'■■  ""•'^<'   '"•  "•■tioM   definitely   associated   with    feediufi   time, 

was  sutlicieut  to  excite  the  secretion.  The  ai)i)etito  .juice  when  started 
was  found  to  persist  for  some  time  after  the  stimulus  causing  it  had 
heen   removed. 

rarlM.n  has  succeeded  in  confirminfj  in  man  most  of  these  olisorva- 
ti.ms.  lie  noted,  however,  that  the  secretion  produced  l>y  scoius;  or 
smellin.;  or  thinking  of  food  is  much  less  than  would  he  expected  from 
Pavlov's  ohservations  on  , loirs.  Kven  when  his  subject  was  hungry, 
•  'arlson  did  not  ohserve  that  the  hriufrius;  of  a  tray  of  savory  food  iiito' 
the  room  caus.Ml  any  secivtion  of  v'astric  juice.  It  is.  of  coui-se,  to  he 
expected  that  the  (piantity  of  the  psychic  secretion  will  lutt  he  the  same 
in  dilTcrcMt  individuals.  It  has  Veen  ohserved,  for  example,  by  Pavlov 
to  var.\  considci'Mbly  in  the  case  of  do^'s,  and  it  is  very  likely  that  it  will 
vary  .still  more  in  man,  with  his  m..r<'  highly  complicated  nervous  system. 
In  no  case  coidd  Carlson  observe  any  secretion  of  <;astrie  juice  pnulueed 
by  havinir  his  patient  chew  on  indilTereiit  substances,  or  by  stimulatinsr 
the  nerve  ...idiii-s  in  the  mouth  by  substances  other  than  "those  dircctlv 
related  to  f(Hid. 

In  man  the  rate  of  secretion  is  i)rop.u-tionaL  to  the  palatability  of  the 
•" 1'  <'"'  smallest  auK.unt,  durinsr  twenty  minutes'  nmsticatiini  of  pal- 
atable food,  beinir  :!i)  c.c.  and  the  larsjcst  V,0  o.e..  in  a  series  of  mfi  obser- 
vations. A  tyjiii-al  curve  showinjr  the  amount  of  the  seerctimi  is  {liven 
in  V]<x.  14!l.  To  construct  this  curve  the  trastric  juice  was  collected  dur- 
intr  five-minute  intervals  while  the  man  was  cliewin<r  a  meal  of  avera<re 
comixisition  and  of  his  own  choice.  An  intercslinf;  feature  depicted  on 
this  curve  is  that  the  secretion  rate  was  hiirliost  in  the  last  five-minute 
period,  this  beinj.'  the  time  during'  which  the  dessert  was  beiiiR  taken, 
for  which  this  man  had  a  (rreat  relish.  Quite  clearly  there  was  a  direct 
relation  between  the  rate  of  the  secretion  of  the  appetite  juice  and  the 
palatability  of  the  food.  Tt  will  further  be  observed  that  it  took  only 
from  fifteen  to  twenty  niiinites  after  discontiiniin<r  the  chewing  before 
the  juico  returned  to  its  original  level. 

The  practical  application  of  these  facts  in  connection  with  the  hvffiene 


I 


I'HVSIOI-OOY   OF   THK   DKiKSTlVK   (il.AXKS  437 

„f  diet  and  the  feeding  of  patients  during  convalescenee,  is  obviously 
very  Rfeat.  However  perfect  in  other  re-ards  a  diet  may  l)e,  it  avi  1 
,„.„i,ablv  fail  t..  be  ditreste.l  at  the  proper  rate  vn.less  it  is  taken  with 
n-lish  Fre.iuent  fee.linfl  with  favorite  morsels  is  more  likely  to  be  tol- 
l.nved  bv  thorough  digesti.m  and  assimilation  than  ..eeasmnal  stuffing 
with  larger  amounts.  We  see  too  in  Iheso  ..xperiments  an  explanation 
„r  the  ^vell-esta!.lished  praetire  of  slMiling  a  meal  with  son  ethmg 
...vory  A  hors  d'onirrr  is  nothing  more  than  a  physioiogie  stimulant 
,!,  appetite.  It  is  also  interesting  from  a  praetieal  standpoint  to  ..bserve 
that  with  those  who  have  a  keen  relish  for  sweetmeats  the  taking  of  des- 
sert has  a  real  phvsi.,lojri,.  signiliean.-e,  for,  as  in  Carlson's  i)atient,  it 
stimulates  towanl  the  end  of  a  meal  a  further  seeretion  of  the  gastrie 


Chewini;  t-xid 


(.arlson.) 

juiee,  and  thus  insures  a  more  ra,.id  .liu'estion  of  the  food.  «!o<..l  eooking. 
it  sh.mld  be  remembered,  is  really  the  fir.st  stage  in  .ligeslion.  and  it  is 
the  only  stage  over  whieh  we  ean  exereise  voluntary  eontrol. 

The  Hormone  Element  in  Gastric  Secretion 

Although  gastrie  dige.stion  is  initiated  by  Ihe  appetite  juiee,  it  is 
.■l.ar  that  this  alone  ean  not  aeeount  for  all  the  seeretion  that  oeeurs 
.luring  normal  gastrie  digestion.  After  an  ordinary  meal  ga..trie  diges- 
tion lasts  usually  about  four  hours,  whereas  we  have  seen,  partieularly 
from  Carlson's  observations,  that  the  appetite  juiee  lasts  only  lor  some 
lifteen  or  twentv  minutes  after  the  exeiting  stimulus  has  been  remove.l. 
The  appetite  juiee,  in  other  words,  serves  only  to  initiate  the  proeess  ot 
seeretion,  and  the  .,"o.stion  arises.  What  A-/,,  w,.  >hr  .rnrl,on  durm, 
the  rc,t  of  gastric  digestion.^    The  answer  was  fur.ashed  by  I'avlov.  who 


[    1 

i 


1 

t 

tH 

;i 

ri 

'  H 

y 

)  ^Pf 

-lis 


Dliil.SllON 


!■ .  '1 


l:i 


iifi!     i 

'ill!  i 

.1 


olps,.,v,.,l  .•niiiMjils  in  «|,i,.|;  ii,.(  only  a  ininiiilnrv  slomaHi  lia.l  1 n  ma.l.' 

I'HI  n  listnla  int„  tl,..  main  slcna.-l.  as  wrll.     Vih-  Ldiaxiur  „f  tl.o  so.-iv- 

'"".'   "'■   "•■"^"•'''   -i"!''''   '1-*   :i    "li «'"iiM   l>c    lollinv..,!   l.v    .•..ilr.-tiiii;   lliat 

ulnrl,  was  s....n.l,.,l  in  tl„.  niinialM.c  ,s1o„,arl,,  for  it  was  sliowii,  in  con- 
1i"l  .■x|Hni,i.,its,  llial  this  scnvlioi,  runs  stri.-tly  j,arall..l  with  that  in 
•  Ik'  iiiiiin  stoma.-ii,  hcin-  (luanlitatix  •■!>  a  .l.linit..  tVaclion  of  it  ar  onl- 
ini:  to  til."  .-..hitiv,.  siz,.  of  th..  minialuiv  stomach  and  .inalitativHv 
I'l'iiticnl.  The  MiinialuiT  stoii,a.-h,  in  other  wonls,  .nirn.rs  tlic  events 
lit  sccrclioii  in  the  main  sloniarii. 

It  «as  ohs,.|v,.,|  tiiat  wli.n  the  animal  was  allow,-.!  to  take  the  f...,.] 
into  th.-  main  stonia.-h  l.y  Ih,-  m.,nth  an.I  .sophai^nis,  the  s.-eieti-m  from 
Iho  mniiatnre  sloma.-ii   .•ontinn.-.j    t..   Il.nv   until    the   pna-ess  of   .rastrie 

-isestmn  ha.l  I,,-.., ,M,.l..t,.,|.  a   .vsnlt   whi.-h   was  .|uite  .lifT.-rent"  fnm, 

that  ohtairn-.l  afl.-r  sham  IV..,|in-.    Tl niv  ,M.ssil,l,.  -xplanation  f.,r  this 

result   ,s  that   Il„.    f 1   in   tin-  sl.,ma-h   s.-(s  up  s.-eretion   as  a   result    .,f 

hn,I  .ln,n,l.,l,„„.     T..   inv.st iv^at.-   th.-  nature   of  this   lo.-al   stimulati.-n 

"'"■""•''  ""''■''•■ ••''  '"•  <-hrw\.n],  f I  an.I  oth.-r  snl.stan.-.-s  were  |,la.-..| 

111  the  man.  ston,a.-h  t1,r.,u-h  the  -astrie  fistula  without  the  aninu.rs 
;'M.wl.-.I-,.  s..  as  to  avoi.l  possii,!,-  j.sy.-hie  slimulati.m,  an.I  the  s,-ereti..n 
"i  .'rve.l  fr.,m  the  miniature  st.mia.-h.  AVIn-n  the  mue..us  m.-ml.rano  of 
tiie  mam  sl.,ma.-h  was  st  innilat..]  nnehani.-ally.  as  l.v  plaein..  inert 
"'M-ets  su.-h  as  a  ,.i.-..e  of  sponyv  ,n-  san.l  in  Ih.-  sl.unaeh,  n..  s.^-reth.n 
"-•-•urr.-,'  Kvi.lently.  ll,eref..r.-.  the  stimulus  is  .|..p,.n.lenl  upon  s.,m.- 
eh.'mi.-al  .piality  of  th.-  f 1. 

,.';•'■  '"t'"'l'i'-iii^'  vari.Mis  f Is  it   was  foun.l  that   th.-re  is  .-onsiderahle 

«lilT.  rene,-  ni  th,-  ,|....mv.-  to  uhi.-h  th,-y  .-an  .-xeit.-  the  s.-er.-tion  Water 
.■u'l-'  white.  hr,-a.l  an-l  star.-h.  ^w.r.-  all  fouml  t.,  hav,-  v.-rv  little  if  anv 
rn-rf.     On  th..  .,th.r  han.l,  u  h,,,  i.rol.in  that  ha.l  heen  partly  ,li-est<-i| 

l.y   means    of   p,.psin    aii-I    hy,|, hl..rie   a.-i.j    was    intr.,.1ue.-,i    int..    the 

st..ma..h.  It  imm..,liat,.ly  ,.a||.-,i  forlh  a  s.-.-r.-tion.  The  eon,.|usi.,n  is  that 
the  partly  .liur,.st,.,l  pr.Mlu.-ls,  .-v.-n  .,f  insipi.l  f.,o.l,  ar,-  .-ajmhle  of  .Hre-tlv 

'■'■''"'""   ""■   ^ •'■"""■      ''"    -^^   iM'-Iii.Ic   prot.-.,s,-s  an.I    |..-pt.,nes    an.l    ii 

uas  th.-ef„re.  „f  n.-.-a,  m   u,   lin.l   that   a   s.,luti..n  of  ..on'imen-ial 

Iiept.m,-  IS  also  an  .-iT-.-tiv.-  s,  nnnlus.  This  is  a  r,-sult  of  ,l.-.-p  si.rnili.-an.-e 
f.u-  It  in.Ii.-at.-s  that  the  fooj  „hi,.|,  has  h.-.-n  partiallv  .li^r,-st,.,l  l.v  the 
iipp.'tite  jui.-.-  will  s.-rve  as  a  slimuhis  to  .•..iitinu.-.l  s.-er.-ti.m. 

The  psyehie  jui.-,-  has  l„-en  aptly  ,.all<-,!  the  "iu'niti.Ml  jui.-.-  "  1,,-eaus.- 
l.y  pro,l„ein!r  partial  -li-.-sti.,.,  it  serv,-s  to  ignite  the  pr.M-ess  of  .rastri,- 
seer,-tion.  Kx,..-rini.ntal  ,vi,|,-n,.,.  of  its  !rr,-at  imiwutanr-e  in  -astri.- 
.Iiir.-sti.m  was  s.-.-ur.-l  l.y  I'avlov  in  esp.-rim.-nts  in  whi-h  ho  pin-,-' 
^ve,3he,i  quantiti.-s  ,.f  meat  atta.-ho.l  to  thr.-a.ls  in  the  st.nnaeh  throu-'h 
a  fiastne  fistula.  an.I  after  s..me  time  r.-m..vo,l  them  and  determine,!  l.v 


i-iiYsi()i.o<iV  (IK  riii:  i>i(ii;sTivi 


.AN1>S 


4:?!) 


tlu>  diffiMTiico  ill  weights  llio  i-xtcnt  to  wlii.-li  llicy  Im.l  l.c-ome  .litjcslcd. 
II  was  found  that  when  tlio  aiipHitc  .jui.-o  was  .-x.-itiMl  l.y  sliam  iVcliii!,' 
Mt  the  sai.io  lin.o  tliat  food  was  plao-d  dir.M-llv  in  tlu>  slomacli,  its  digos- 
tioii  was  imi.-h  inoiv  rapid  tlian  in  cases  in  wl.i.-li  it  was  plar.-d  111  the 
stomach  without  the  aniiiiai's  kiiowins.',  as  when  ho  was  asleep. 

Other  foods  haviiijr  a  direct  stimulatins.'  elTect  on  the  gastric  secre- 
ti„„  are  meat  extracts  ami,  to  a  certain  extent,  inilU.  This  elT...-t  of  meat 
..xtract  is  interestinst  in  connection  with  the  practice  of  takinj:  soup  as 
a  first  or  early  stajre  in  dininir.  It  not  only  excites  the  ap|>etitc  juice, 
i„il  also  serves  as  a  direct  stimulus  to  the  gastric  secretion. 

As  to  thr  uatiirr  of  the  mrrhoiiism  h>i  whuh  this  (llrrrt  srrntlon  tnhrs 
,,l„rr.  it  was  shown  by  ropiclski'"'  that  the  secretion  still  occurs  after  all 
,1„.  nerves  i.rocee.linsr  to  the  stomach  are  cut.  Evidently,  therefore,  it 
is  independent  of  the  extrinsic  nerve  snpi.ly  of  the  viscus.  As  a  result 
of  hi>:  experiments  ropi<'l^l<'  concluded  that  th  •  secretion  must  (h'I.end 
on  a  local  rellex  me<liated  through  the  ner\e  structures  present  in  the 
walls  of  the  stomach  itself.  Another  explanation  of  the  result  has, 
hcnvever,  in  recent  years  l.een  given  more  cre.lcn.'c  l.y  the  experiments  oi 
r.iyliss  and  Starling  on  the  inlluence  of  hormones  on  the  si'cretion  ot 
,,;„,,,,„i,   j„i,,.    |,r.    page    4-j:.l.     Kdkins-    .suggcstc.l     that     a     similar 

process   in  tlie  stomach   might    ac nut    for   the   continued   secretion    ol 

•Mstric  iuice.    To  test  the  possibility  this  investigator,  after  ligatuig  the 
rardiac    sphincter    in    anestlietizcd    animals,    inserted    a    tul.e    into    the 
pvloric  end  of  the  stomach,  thn.ngh   which   he   pla.-ed   in   the  st.miach 
iilM.ut  r.O  c.c.  of  i.liysiologic  saline.     After  this  had  l.ceii  in  the  stomach 
for  an  hour,  he  found  that  no  water  was  al.s.n-l.ed,  and  thiit  if  the  tlui.l 
«,is  removed  aft.u-  this  time,  it  contained  neither  hydrochhnic  aci<l  nor 
pepsin.     On  the  other  hand,  if  during  the  time  the  saline   was  in   the 
stomach  a  decoction  of  the  mucous  meml.rane  of  the  pyloric  end,  made 
either  with  peptone  solution  or  with  a  solution  of  dextrine,  was  injected 
intravenously  in  small  ((uantities  every  few  minutes,  it  was  found  that 
the  sMl-lie  contained  distinct   .|'i:i"t'''"''<  "»"  hydrochloric  acid   and   pepsin. 
Furthermore,  it  was  found  that,  if  the  pei.tone  .s..lution  or  the  dextrine 
soliiti(m  alone  was  injected  intravenously,  there  was  n..  such  evidem-e 
of  gastric  secretion.    The  conclusion  which  Kdkins  drew  from  his  experi- 
ments is  to  the  ofTect  that  the  half-digest e,l  produ.-ts  of  the  earlier  stages 
of  gastric  digestion  act  cm  the  mucous  iiicmlirano  of  the  stomach  so  as  to 
pnuluce  a  hormone,  which   is  then  carried  hy  the  hlood  to  the  cells  of 
itie  gastric  glands,  upon   which,  like  secretin,   it   directly   develops  an 
cN.-ithig  efToct.     Tliis  hormone  has  been   called  qnsfrin.     These  observa- 
tions of  Edkins  have  been  confirmed,  and  they  explain  very  ijimply  how 
gastric  .secretion  is  maintained  after  the  cessation  of  the  secretion  of  the 


;  ■.!  i. 


\l 


• 

■  . 

• 

'  ,1 

!«  ^   i; 


i  1 


1  , 

i  it 

itt 


440 


DIGESTION 


appetite  jiiico.'"  ]\\  such  a  inpchaiiisni  gastric  juice  would  continue  to  Iv 
secreted  so  loiiij  as  any  half-jllKested  food  remains  in  the  stomach. 

The  action  of  srastrin  is  tlie  first  instance  of  a  hormone  control  of  th( 
difrestive  fila.Mls.  In  the  earlier  stages  of  diKfstion,  the  secretion  of  saliva 
and  apiiclilc  juice  is  mediated  through  the  nervous  system,  hecause  these 

juices  must  he  produ 1  promptly.     In  the  later  stages  of  uastrie  difjes. 

lion,  such  pioMiptitude  in  response  on  the  i)art  of  the  fjlaiul  is  no  longer 
necessary,  so  that  the  slower,  more  continuous  process  of  hormone  con- 
trol is  suflicient. 

Quantity  of  Gastric  Juice  Secreted 

Accordiiisr  to  Carlson,  the  totjd  amount  of  gastric  juice  secreted  in 
man  on  an  average  meal  comp(.sed  of  meat,  I.read,  vefjetables,  cofTee  or 


I'l.sli.  Jiriluii.. 


ISn-aii,  Jon  ^in. 


.Milk,  (.110  c.c. 


l-'iR.    I^rt.      I'ul.u- 


.ntinut.r.s  of  KaMric  i„i«-  Mirrt.-,!  after  .11.  Is  .,f  tikmi,  brca.l,  and  nnlk.      (Fr,. 
Pavlov. ) 


milk,  and  dessert,  amounts  to  about  700  c.c,  beiiiff  divided  into  200  c.c. 
in  the  first  hour,  l.'JO  in  the  second,  and  :?oO  c.c.  during  the  third,  fourth 
and  fifth  hour.s.     These  fljiures  were  estimated  partly  on  the  basis  of 
oiiservatioiis  made  on  the  man  with  the  gastric  fistula,  and  partly  from 
the  datfl  supplied  by  i  i.vlov's  observations  on  dojrs.     Carlson  believes 
that  I'avlov  overestimated  the  relative  importance  of  the  appetite  juice 
11.  -astnc  digestion.     He  found,  for  example,  that  after  division  of  both 
vagus  nerves  in  dogs  normal  gastric  digestion  might  be  regained  a  few 
days  after  the  operation,  although,  of  course,  under  such  circumstances  no 
appetite  juice  could  have  been  secreted.    Moreover,  he  ob.served  that  caf; 
when  forcibly  f,.d  with  unpalatable  f.iod  may  digest  that  food  as  rapidh 
as  when   they  eat   voluntarily.     In  support  of  his  contention,   Carlson 
states  that  he  has  frequently  removed  all  of  the  appetite  juice  from  his 
pati.        s  stomach   before  the  masticated  meal  was  put  into  if   without 
any  evident  interference  with  the  digestive  process. 
Fat  has  a  distinct  inhibiting  influence  on  the  direct  secretion  of  gas- 


\\\ 


rf 


,!1 


PHYSIOLOGY   OP   THE   DIOESTIVK  OLANOS 


441 


trio  juice;  croam  takes  considerably  longer  to  be  be  digested  than  milk, 
iind  the  presence  of  oil  in  the  stomach  delays  the  secretion  of  juice  poured 
out  on  a  subseciuent  meal  of  othernise  readily  digestible  food.  By  col- 
lecting all  of  the  gastric  juice  from  the  miniature  stomach  after  feeding 
l.y  mouth  with  (juantities  of  different  protein-rich  foods  containing  the 
same  quantities  of  nitrogen,  interesting  observations  have  been  recorded 
concerning  the  amount  of  juice  secreted  and  its  proteolytic  power.  The 
results  of  some  of  the  cxpcrimcnis  arc  shown  in  the  accompanying 
curves  (Figs.  150  and  151). 

It  will  be  seen  that  the  most  abundant  secretion  occurs  with  meat,  that 
of  milk  being  not  only  smaller  but  also  slower  in  starting.  The  digestive 
])ower  is  greatest  in  the  case  of  bread. 


Hours  (23458782343e78    9    2    3    ♦  5    « 


III.  ail,  J(IO  Km. 


Milk,  (.(il)c.c. 


IMcsll,  JOO  Kill. 

I-,K     151       l)iK.-.liv.-  l.nwer  of  tlio  jui.-.-.   .t   nl,aMir<-,l   l.y   Ih.-  IcnKth   nf   tlir   ,.rol.in   ciniml    .liK.sl..l 
in    Mrtt's    tillKs,    with    ili.ts    if    fli-sli,    lircul.    ami    milk.       (l-ronl    lavlnv.) 

THE  INTESTINAL  SECRETIONS 

Pancreatic  Juice 

Regarding  the  natural  secretion  of  pancreatic  juice,  little  need  be  added 
to  what  has  already  been  said  (sec  page  42.")).  The  secretion  lH'!,'iiis  wlicii  the 
chyme  enters  the  duodenum,  and  attains  its  maximum  when  the  outflow 
of  this  is  greatest.  By  collecting  the  juice  from  a  iteriiianent  fistula  of  the 
l)ancreatic  duct,  it  has  been  found  that  the  amount  varies  with  different 
foods.  When  quantities  of  food  cont.iining  equivalent  amounts  of  nitro- 
'_'en  are  fed,  the  greatest  secretion  is  said  to  occur  with  bread  and  the  Ica.st 
with  milk.  Such  differences  are  probably  dependent  upon  the  amount  of 
iicid  secretwl  in  the  stomach  and  passed  on  into  the  duodenum.  It  was 
llu.iight  at  one  time  that,  besides  variation  in  quantity,  the  nature  of  the 
enzymes  in  the  pancreatic  juice  might  vary  according  to  the  kind  of 
food.    This,  however,  has  been  shown  not  to  be  the  case. 


a 


Mi 


M 


1 

»  ^' 

1 

ft  1 

1 

i  1 

1 
i 

!k^ 

1 

||; 

I 

1  ^ 

i 

i 

mmii 

Uil 

Wi 

442 


KKlKSriON 


I  (        .! 


4  ■ 


BUe 

The  MTicli.iii  of  l.il,.  runs  i)i-;iclically  paiallfl  with  tlial  of  pancreatic 
Juice.  Tlic  liver  is  pr«nliicin«>:  liile  more  or  less  contimioiisly,  since  l)esi(lcs 
i'eiii!,'  a  iii(;esti\c  lluid  it  is  alsii  an  excreloi'y  product.  The  liiie  produce.l 
between  the  periocls  of  (|ii;esti(in  is  niainlv  stored  in  tlie  -.'all  Madder. 
When  the  acid  chvine  eonies  in  contact  with  the  duodenal  iiincous  meni- 
hraiie,  it  excites  alTcrent  nerve  endin!.'s  that  cause  a  reliex  contraction  oi" 
till'  srall  hladder.  and  this  expresses  some  ot'  the  hile  into  the  duodenum. 
The  secretin,  which  the  acid  at  tlii".  .sanu'  time  produces,  besides  nlTocfinf; 
tlic  iianciras.  acts  on  the  liver  cells,  stimulating'  them  to  the  increased 
secretion  of  hile.  Thus,  l.y  a  nervous  reflex  operating'  on  the  fjall  bhuhh'r 
and  later  hy  a  hormone  mechanism  operatiuf.'  on  the  liver  cell,  tlie  iiu-reased 
si'creliim  of  bile  is  insured  tIiron«:houl  ditfestion.  Of  the  bile  diseharifcd 
into  the  intestine,  ii  certain  i)ropi.rtion  of  the  bile  salts  is  reabsorbed  into 
the  portal  blood.  When  these  arrive  at  the  liver  they  also  excite  .secre- 
tion of  bile,  thus  assislin<;  .secretin  in  maintaininj;  the  secretion  tlirouj;h- 
out  the  process  of  intestinal  di-jestion. 


II     !• 


'■„■    . 


■M'>^ 


'■'"■    '''       '•""''   ■■>"'<'•■■'■<■    'Xur    IMMU   „tr  ,\w   i...,i,.,„^.   ,„l.ii,K   til.    ,Hiv,>   niMMi.iK   I.,   tlir    n.i.MI. 
l-.n,.,,,,   ;„„i    r.  innuim    il„-    I,,..,,    i.,    ,1„    .-,l„l,,m,n    f..r    -.uw    linu-.      (l-„„n    Jarki-nn.) 

Intestinal  Juice 

The  secretion  of  intestinal  juice,  or  succus  enterieus,  can  obviously  he 
.studied  only  after  isolatintr  portions  of  the  intestine  and  eonneetiufr  thctn 
with  fistuhe  of  the  abdominal  walls.  It  a))pears  here  aftain  that  both  a 
nervous  and  a  hormone  mechanism  exist.  ]\reehanical  stimulation  of  tin 
intestnial  mucous  membrane  causes  an  immediate  outflow  of  intestinal 
.jnieo.  the  purjx.se  of  which  inider  normal  conditions  is  evidently  to  assist 
in  moviii};  forward  the  bowel  contents.  This  mechanically  exeited  .juice 
does  not  contain  any  enlerokinnsr  and  only  small  amounts  of  the  other 
enzymes.  Further  evidence  for  nervous  c(mtrol  of  the  secretion  of  intes- 
tinal juieo  has  been  obtained  by  isolating  three  pouches  of  intestine  he 


i'iivsi()i.(MiV  (IP  Tin;  KKiisTivi;  (iianhs 


44:» 


twc.'ii  limituiTs,  ami  tlicii  .Ifii.Tvafmu'  llif  ci'iilnil  i.oucli  l.y  caicfully 
.•iillintf  all  the  iicivos  witlmul  \voiiii«liii«  tlio  1)1oimI  vessels.  On  ivturniii« 
the  ix.iU'lu's  to  flic  alxlonieii  aii.l  lcaviii«;  tlwin  srvcial  limirs,  it  lias  been 

l',,iiMil  that  tliP  Iiiiihllc  li'iiicli  l>(< es  (listciiil.Ml  with  sfcicli.iii,  whereas 

the  two  eii.l  pouehes  leiiiaili  emiitv  (  Tiif.  l.')2).  If  the  jioiiehes  are  left  for 
several  ilnvs  in  the  alxloineii,  houev.'r,  the  seeretioii  fn-m  the  deiiervated 

tioii  disappears  asiain.     The  ..xiilaiiatioii  of  the  result  is  possil.ly  that 

the  nerves  under  ordinary  i litions  eonvey  impulses  to  the  intestinal 

.^'lands,  whieh  lonieally  inhiliit  their  aetivity. 

The  existeiiee  of  hormone  eontrol  is  evideneed  liy  the  faet  that  mi 
niterokinase  is  present  in  the  intestinal  juiee  unless  panereatie  juiee  is 
placed  in  contact  with  the  mueous  memlMane.  Injection  of  pancreatic 
iuico  into  the  Mood,  however,  does  not  cause  any  se.'retii.u  of  intestinal 


,1 


juiee;  whereas 


hei 


the  inject ifin  of  secretin  has  such  an  effect. 


i  t 


!',    l^ 


i'(i    " 


•  IIAI'TKU  M 

TMK  MKCHAN'ISMS  OF  DICKSTlOX 

MASTICATION.  DEOLUTITION,  VOMITING 

Mastication 

I'.y  thr  iiMivciiicMfs  of  ilu-  lower  jaw  mi  llic  upper,  the  two  rows  ol 
teelli  eoiiie  tojretljer  so  iis  to  serve  for  liifiiiu'  or  eruslini.„'  llie  food.  TIk 
resiilfiin;  ('01111111111111011  of  the  foo.1  forms  the  lirst  step  in  tlici'stion.  Th. 
up  ami  (h)Wii  niotioii  of  tlie  lower  jaw  results  in  liilini,'  hy  the  iiieisors, 

and  after  the  mouthful  has  I.eeii  takfii.  the  side  to  side  vemeiits  eiial.l.' 

tlie  Krindiiiir  teeth  to  eriisli  and  hreak  it  up  into  fraj;ments  of  the  in'(i|Mf 
size  for  swallowlnj.'.  The  most  suHal)le  size  of  the  mouthful  is  alioiii 
T)  i-.c,  hut  lliis  vari.'s  j;reatly  with  lialilt.  AfKi  mastieation,  the  iiiaNs 
Meinhs  from  :{.2  to  (i.')  jrm.,  al)'>ut  one-fourth  of  this  weight  heiii",'  due  1.. 
saliva.  Tlio  food  is  now  a  semilluid  mush  eontainin};  particles  whldi 
are  usually  less  than  2  mm.  in  diameter.  Some,  however,  ma.s  measiii. 
7  or  even  12  mm. 

Determination  of  the  jnoper  de-;ree  of  lineness  of  the  food  is  a  fiiih- 
tioii  of  the  tonsjue.  j.'ums,  and  eheeks,  for  wliieli  imrpo.se  the  miieoii- 
mei;)l  tane  eoverinjr  them  is  supplied  with  very  sensitive  toueh  nerv. 
('ndln-.'s  (see  paj;e  7!>4,.  The  sensitiveness  of  the  tonjrile,  ete.,  in  tliiv 
roKard  explains  wliy  an  ohjeet  whieh  ean  seareely  bp  felt  by  the  (iiisrci- 

seems  to  he  quite  lar<re  in  the  nth.     If  some  i)artieles  of  food  that  ai. 

too  lar-re  for  swallowlntr  ha|)pen  to  l.e  eariied  liaekward  in  the  monil. 
the  toiifiuo  returns  them  for  further  mastieation. 

The  saliva  assists  in  mastieation  in  several  ways:  (1)  ],y  dissolviii' 
s(mio  of  thp  food  eoiistituents:  (21  by  iiartly  diifpstiii};  .some  of  tli 
starch:  (.'{)  by  .softeninjr  the  mass  id"  foo<l  so  that  it  is  more  readil 
prushed:  (4)  by  eoverinsr  the  bolus  with  mupus  so  as  to  make  it  mm 
readily  transferable  from  place  to  place.  The  secretion  of  saliva  1 
tliprefore  stimulated  by  the  chewin<r  movemPiit.s,  and  its  pompositi(  ■ 
varies  according'  to  th.'  nature  of  the  f(MKl  (  pafre  4:!!).  In  some  aniiiKiN 
such  as  the  cat  and  df)<r,  mastication  is  unimportant,  eoatin}.'  of  the  food  will 
saliva  bein;.'  the  only  cbaiisre  which  it  uiidoi'ioes  ill  flip  mouth.  In  tiiii 
the  ability  thus  to  bolt  thp  food  can  readily  be  apcpiirpd,  not,  however 
without  .<iomp  dptrimput  to  thp  pfficipiu-y  of  digpstion  as  a  Avholp.     Sn!'^ 

444 


Tin:    MIlIIANlSMR  OF    IMOKSTIOV 


448 


stiircliy  foo.1  is  littl.-  cIhwo.I,  tin-  I'liu'lli  of  tiiii.-  miiiin',1  for  \\\v  luaHti.-H- 
tii.ii  of  otluT   fo.Mls  (U-iHMi.liiiif  iiiaiiily   on   thoir  iiatuii'.  luit   nlno  to  a 

If  cnn  not  1»P  too  stioiiuly  iiiHisfcd  vip<iii  that  tlip  act  of  mnsticution  is 
,,r  far  more  import aiioi-  tliaii  iii<T«>Iy  to  liivak  up  aii.l  prepare  the  food 
for  swnllowiiit'.  It  causes  the  food  to  l.c  moved  al>oiit  in  tlic  mouth  so  M 
I„  develop  its  full  olTect  (.u  the  taste  l.uds:  the  ciushiint  also  rolcasM 
„dois  which  stimulate  the  olfactory  epithelium  (»u  these  stimuli  dopend 
the  sntlsfnction  and  pleasure  of  eatinir.  which  in  turn  initiate  the  proeoss 
.if  irastric  dis:estion  (see  pau'c  4:>.'>>. 

The  benefit  to  di^'estion  as  a  whole  of  a  lariro  secretion  of  saliva,  brouRht 
ahoHt  hy  persistent  ehewinc  has  l.een  assumed  l>y  sorre  to  ho  much 
..'renter  than  it  really  is.  and  there  has  existed,  and  indeed  may  still 
.  xist,  a  school  of  fa.ldists  who,  l.y  delilierafely  chewinR  far  beyond 
the  necessary  time,  imaiiine  themselves  to  thrive  better  on  less  food  than 
those  who  occupy  their  time  with  more  profitable  pursuits. 

Defrlntition  or  Swallowing: 

After  beiti?  masticated  the  food  is  rolled  up  into  a  bolus  by  tb<^  action 
„(  the  tongue  asrainst  the  pa'ate,  and  after  beini;  lubricated  1  .  iva  is 
iniived.  by  elevation  of  the  front  of  the  top.irue,  towards  the  ba.  of  the 
iiK.ulli.  This  constitutes  the  prsi  slaf/r  of  swallow! nsr,  and  is.  so  far,  a 
\. limitary  act.  Abotit  this  time  a  sliirht  inspiratory  contraction  of  the 
diaphrajnu  occurs  the  so-called  respiration  of  swallow iiiff— and  the 
„ivJohvoid  ,\\uMy  contracts,  with  the  eonse<iuence  that  the  bolus  passes 
iHlween  the  pillars  of  the  fauces.  This  marks  the  beitiniiinff  of  the 
^.,ond  sh„f,  the  first  event  of  which  is  that  the  bolus,  by  stimnlatins: 
Miisory  iH-rve  endin-.'s.  acts  on  nerve  centers  sittiated  in  the  medulla 
uMonirata  so  as  to  cause  a  co.inlinated  series  of  movements  of  the 
iMiisdes  of  the  pharynx  and  larynx  and  an  inhibit  ion  for  a  moment  of 
llii'    respiratory   center    <  pajre   :>:!"J). 

The  movements  alter  the  shape  of  the  i>liMrynx  and  of  the  various 
i.peninffs  into  it  in  such  a  manner  as  to  compel  the  bolus  of  food  to  pass 
into  the  esophajius  (see  Fi-  irvD:  thus.  (Pi  the  soft  palate  becomes 
rlcvated  and  the  posterior  wall  of  the  pharynx  Iniliics  forward  so  as  to 
shut  off  the  posterior  nares,  (2)  the  posterior  pillars  of  the  fauces  ap- 
I.n.ximate  so  as  to  shut  off  the  mouth  cavity,  and  i'^)  m  about  a  tenth  of 
;i  second  after  the  mylohyoid  lias  contracted,  the  larynx  is  pulled  up- 
wards and  forwards  under  the  root  of  the  tongue,  which  by  beins 
'Irawn  backwards  becomes  banked  up  over  the  laryngeal  opcnitig.  This 
pulling  up  of  the  larynx  brings  the  opening  into  it  near  to  the  lower 
!  nif  of  the  dorsal  side  of  the  epiglottis,  but  the  upper  half  of  this  strue- 


'Hi 


„   .   1 


1: 

i 

1 

1   ' 

li  1 

;    A 

u 

: 

p 

m 

44A 


liliil  >TH»N 


tiiiT  |ir(i.|ri'ts  l.i'MiHcl  iiikI  ««»r\is  us  ii  IciIkc  to  uiiii|i>  the  liulii.t  sal'i'ly  piiNi 
tliis  i-iiii.-,il  |iiiit  i.r  its  ccim-sr  iti  As  a  fiirtlicr  siifcuiinnl  atrjiiiist  imi\ 
fiili-;  III'  r.Pdil  iiitii  the  .lir  pMss.itris,  the  hir.Mi'.'fiil  ..|n'iiiiiu  is  narrownl  l.< 
a|i|ir.i\iiiijiti..ii  of  tlw  Iriir  iiml  tlir  I'lilsc  \i.cal  i'i>i«|s. 

So   l;ii'   till'   fiilrr  wliicli    |i|'ci|ii'|s  llic  licihis   is  inailily   tlic  coiit  rai'tioli   <<( 
llir    lii\  liiliMiiil,   assisfi'il    li\    th<>    lrici\ciii.-iits   nl'   the    iniil    of   tile    fiiiiu'lH' 
Wliiii   it   lias  nai'lii'.l  llic  I.iwit  tml  id'  tlic  pliarv  ii\,  linwcvcr.  the  Ixilii^ 
irailily    lails   iiitii  tin'  isii|iluii;ils.   which    has  Imcdiuc  (lilatnl   on   nccoiiiit 
111'  a  icIlcN   iiiliiliitioii  III'  the  ciiiisliiclor  llMlsclcs  of  its  ll|i|icr  finl.     This  sn 

•'•'il''''!   "*' ii>l   stai!i>  111"  suall.iw  ill!.'   is,  f  licnl'iirc,  a   ciiinplrx    ciiiirdiirati'.l 

iiiHMiiHiil  iiiitiati'cj  l.y  all'iiciit  stiiiiiili  ami  iiiNolviiiu'  nripi'o.-al  notion 
111'  \ariciiis  i;i,iii|is  ..;■  niUNrji.s:   iiihiliit''iii  .i|'  tlir   ifs|iiiatciry  iiiii.si>lt>s  ami 


'■■"■■■      '"'  ■','"       ■'•■■"f-,     "I"     ''      '••'•'■      !''"'■      I"      «"       '■'•'U-V      ..(      II:,-      t      nf      ||„.      |„„Wllr.      tllr      -,.11 

r.ll.,1.  .    ill,     >|.n:l..|',,    .,,„!    Il„-    l.,,>,iv    ,|,srn,i;    ll,.     , .|    ,i.,i.-.     ..t    >«  ,,ll,,«  inn^      Tli.-    llii.k    .|.,lt..|    In. 

Ii..,i    .I'lrmii    ■.„.,||.,,i,ii,; 


l,'.li'.,l,  -    III, 


of  thiiM'  that  I'l.nstiii-t  thi'  iMi|i|iairus,  aiitl  stiiiiiilatinii  ol"  fhnsi'  tlinl 
I'li'vatr  tlic  palate,  the  lunt  nl'  till'  liiiivriH',  ;iii<l  thr  l.'ii'yiix.  If  is  pui'iK 
ail   iiiMiliiiitarx    piociss. 

Til.'  //((/•(/  .s7i/,/i    „/'  ,l,,iliilitniii  I sists  ill  till'  passai.'!'  of  tlii>  swallouc.l 

'' '  "I""'-'  "!•'  <'sopliay:iis.     Tlii'   iiii'.'haiiisiii   liy   whii-h  this   is  ilmii'  <li- 

pfinls  \i-\-\    iiiiii'l till'  physii-al  rniisisli'iii-i'  of  tlii'   t'lioil.     A  solid  biilif< 

that  iiii.ic  111-  li'ss  tills  till-  isophatriis  (Xi-ili-s  a  typii-al  ])i'iislaltii'  \\a\r. 

wliii'h  is  chiiiai'li'ii/iil  Ky  a  ililafat i.ni  nt'  the  csiiplia-jiis  i liati'ly  ii. 

I'idht  i.j"  au'l  a  <'M!!stii..ii,,!,  ,,vc!'  :ii!i|  lii'liiml  thr  l-.nhi-.  This  ^\  avt- lia'.f'- 
(li.uii  till'  I'siiphaLMis  ill  man  at  siu-h  a  rati"  that  It  frai'lii's  the  rardiar 
sphiii.-ti'i-   ill   ;ilHiiit    live  ur  six   si'i'iiiiils.     On   arriviii';   licru   the   i-anlia'' 


TIIK.    MKIIUMXMS   DV    l>Ui>;HTIIlN 


447 


sphiiu'tiT,  or 


rilinaiilv  roiitiai'lcl,  i«'lii\«'s  fi.r  a  iii<.iii,nl  m>  tliiil  llic  Ik'Iiih 


luiHM'M  into  till'  Htiiniiii-li.     Ill  iiiaiiv  iiiiinin 

tlic  )nTiMlnlti('  wave  Inivils  iiiurli  iimn-  riiimllx 


Is,  iiicluilini;  niiiii  Mini  the  cut, 


ipluiHiii*  tliK"   loW"!"  "lo^Mt   I'lcail^t 


ill  llif  iipjicr  |>ait  of  lilt' 
f  ililTcrr s  ill  tlif  iiatiirc  of  tin* 


iiiiisciil'ir  coat,  this  hriiiu  of  (In-  stiiati'd  vjirifty  aLovr.  aii.l  "f  llif  iioii- 
viiial<'<l  lii'low.  Til.'  |iiir|iosc  of  more  lapiil  iiiovrmint  in  llir  iipiwr  part 
IS  no  iloiilil  tli/il  llif  I'oIm-*  iiinv  1h-  liiininl  past  flw  it«:i,.iis  w  hen-,  l>y 
,lislrii.rniir  tlic  csopluiuus,  il  ini«lit  iiiti-rlViv  uilli  llu>  fmiHion  of  n.-iirl'- 
l.uiiiiur  sliurtiin's,  .^n.-li  as  tlw  lu^ait.  I.i  otlxr  aiiiiiiaU,  as  iIh-  .l"if.  tli<- 
iMiisiMilar  lilxT  is  sliiatr.l  all  alonu'  tli<'  csopliairus.  nml  tin-  Ih-Iiis  of  foo.l 

ivspoiitlinulv  tnivfis  at  a  iiiiifoiiii,  'inick  rali>  all  tin-  way.     Il  lakos 

Miily  about   four  st Is  for  llir  liolus  t..  n'a<-li  llu'  stoinarli  in  tlu-  «loir. 

Thr  pcristalli.-  «a\.'  of  tin-  upper  part  of  tin-,  csopliauus  in  tlir  <at  ami 
luvsiiinaUly  in  inaii.  niilik.'  Ilial  of  tin-  iiil.-stiiifK  Is,.,  pai;.'  4titil.  is  Hani 
inittc.l  l>y  tilt'  osopliaKoal  l.raiiclirs  of  flu-  vauns  ii.rvts.     If  tli.'si'  an- 

MViic.l,  l.iit  111.'  iiiiiMMilar  ooats  left  iiilact.  the  csopliatrns  I oincs  dilati"! 

:,l«.\..   til.'   I.'V.-I   ..f   111"  s..,'    .r   MM.I   .•ontnirh.l   li.lovs.   aii.l    no   p.^ristalti.- 
uiivc  cnn  pass  nlmiu'  it:  oi      in-  otln'r  liaiol,  llif  niiisi'iilar  coat  may  I'C 

..v.ir.l    (l.y   c.usiruii:,   etc.,    I.nt    tlic    pciislallic    \vavc    will    ( tinuc   to 

imvi'l,  jtroviilcil  no  ihiiiiairc  has  liccii  ilciic  lo  the  n.-rvcs. 

Ill  the  lower  part  ..f  the  .■sophau'iw.  however,  the  wave  of  peristalsis, 
like  ihnt  of  the  inlcstines,  travels  in.lepcn.lentl\  of  extrinsic  nerv.  s. 
Tiiis  has  been  observed  in  animals  in  which  all  of  llie  extrinsic  iiervrs 
liMve  been  cut  some  time  ].revioiisly.  This  .lilTerence  behvecn  the  upper 
Ml. I  the  lower  p..rti..ns  is  associate.l  with  Ilie  .lilTereii.-e  in  Ihe  nature  of 
ilic  muscular  libers  ab.ive  noted  (Melt/.er)." 
The  i.r.)i.ai;ati..ii  .'f  the  wav.-  by  Ihe  nerves  in  the  ujipcr  part  of  the 

.  so|.hai:us  inili.-at.'s  that  the  s ml  slai.'.'  aii.l  the  (iiM  imrt  ..f  the  third 

siM-e  of  .lc.,'lutiti.iii  must  be  reliearse.l.  as  it  were,  ill  the  me.liillary 
.•.HUMS  from  whi.-h  arise  the  nerve  libers  to  the  pharynx  and  the  ufiper 
l.v.is  ..f  the  esoi.hai;iis.  It  is  1houj;ht  that  the  discharges  fron  theso 
l.Ki,l  cctiters  are  controlle.l  by  a  hi'^'lier  Htnilhurhtij  cnihr  sitiiati-d  in  the 
.ii..|iilla  .just  ab.ive  that  ..f  r.'si.ira1i..n,  the  alTerent  stimuli  to  whi<-h 
Pi.i,.,.cd  from  the  i.haiyiix  by  the  fifth,  superior  laryntreal,  and  vai-'us 
iiirves.  The  exact  loeati.in  of  the  sensory  areas  whose  stimulati.m  is 
Miost  effective  in  initiatinir  the  swalhiwin..'  retlex  vari.'s  consi.lerably 
in  .lirtVrent  animals.  In  man  it  is  i.robably  at  the  entrance  to  the 
phaiyiix:  in  the  dot:  it  is  on  the  jxistcrior  wall.  A  f..reisrn  b.xly  place.l 
Miic.'tly  in  the  upper  p.U'tion  of  the  cs..i)lia!rus  of  man  has  been  obscrve.l 
'■■  remain  stationary  until  the  in.li'idual  ma-h'  a  swallowiiK^'  movement. 
The  afforent  fibers  in  the  j;losso|)haryiineal  nerve  exercise  a  i>owcrfnl 
'nhihitorn  influence  on  the  de<;lutitioii  center  as  well  as  .m  that  of  respira- 


hn-iMt 


1?  a 


U  ^1 


448 


nU'.KSTION 


% 


i\ 


If  r 


I  ! 


ii 


lion.  Tims,  if  swallowiiif?  inovcinciits  aic  excited  hy  sliiiiiilatiiij;  Iho  eeii 
tral  »Mi(l  of  tlio  siiporior  laryiifrcal  iierve,  tiiey  eaii  be  instantly  inhibite.l 
by  sininltaneously  stiiiiulaf in-;  the  H;loss<)|)liaryii<;eal,  and  the  i't'spiratiii'\ 
inovompiits  stop  in  wliatevcr  jxisition  tliey  may  have  i)ei'n  at  the  time 
When  the  };loss(>])liai'yns;eal  nerves  are  cut.  tiie  esophagus  cntprs  into  k 
eondition  of  tonic  contraction,  whicii  may  last  a  day  or  so.  This  shows 
that  the  iidiibitoi-y  imi)iilses  are  tonic  in  natnre. 

This  iidiibition  of  the  esophagus  is  indeed  a  most  important  jiart  I'l 
the  proccs.-,  uhen  li<iiti<l  or  si  iiiiliqiiid  fond  is  swallowed.  By  tlie  c<tiitrac 
tion  of  the  mylohyoid  nni.sclc,  llnids  are  (piickly  shot  down  the  di.sten(hil 
esoidiasriis,  at  the  lower  end  of  which,  on  acconnt  of  the  closure  of  tlir 
eai'diac  si)hincter,  they  accumulate  until  the  arrival  of  the  peristaltic 
wave  whii-h  has  meanwhile  been  set  up  by  stimulation  of  the  pharynx 
If  the  swallowiufr  is  immediately  rei)eateil,  as  is  usually  the  case  in 
<lrinkin!.',  the  esophji-rus  i-emains  dilated  because  ])eristalsis  is  iidiibited. 
and  the  fluid  lies  outside  the  caidiac  oiilice  until  the  last  mouthful  has 
been  taken. 

The  Cardiac  Sphincter 

The  passav:e  between  the  esoiiha-rus  and  the  stomach  is  j.'narded  b,\ 
the  eaidiae  sphincter  oi'  car.lia.  This  exists  in  a  permanently  con 
traeted  state,  or  tonus,  sui>erimposed  on  which  from  time  to  time  an 
rhythmic  alternations  of  contraction  and  relaxation.  This  tonus  is  lu'vci 
very  pronounced.  In  man  it  is  said  that  a  water  pressure  of  from  2  to  7 
cm.  applii'd  to  the  esoplijiireal  side  of  the  sphincter  will  drive  air  oi 
water  into  ihe  st«uiin<'h.  this  ))ressuie  beinjr  less  than  that  of  a  column 
of  (luid  tiliinsr  the  thoracic  esoi)h;if;us  in  the  erect  position.  Durin'j 
repeated  de<.'lutition  the  tonus  becomes  less  and  le.ss  marked,  and  aftci 
a  number  of  swallows  the  sphincter  may  become  eomi)letely  relaxeij 
When  this  relaxation  disapjx'ars,  howe\er.  the  spliincter  l>eeomes  mon 
contracted  tlian  usual  and  i-emains  so  for  a  lonjrer  time. 

The  tonic  condition  of  the  sphinctei'  is  controlled  by  the  vas;us  nerve, 
stimulation  of  which  causes  iclaxatioii  with  an  after-effect  of  stron'j 
contraction.  Mechanical  or  chemical  stimulation  of  tlie  lower  end  of  tin 
esopha<rus  inci-ea.ses  the  tonus  of  the  sphincter.  Forein<»  of  the  sphinctt  ; 
from  the  stomach  side  reipiires  a  lii^dier  pressure  than  from  the  esojih.i 
^eal.  Kructatioii  of  <;as,  for  example,  does  not  take  place  until  intr.i 
jrastric  pressure  has  risen  to  about  I'l  <'m.  of  watei'.  In  deep  anesthesia! 
however,  intrnfjastric  i)ressure  may  rise  considerably  hiffher  withon' 
foreinsr  the  spliincter. 

In  ani'nnls  fed  with  slarcli  paste  imiircfrnated  with  subnitrato  of  bi- 
muth  and  then  examined  by  means  of  the  x-rays,  the  variation  in  deirri  ■ 


Tin:    Mirii\NlsM> 


liKJI  STlllV 


44!) 


uT  tout'  of  t  lie  spliiiicirr  Iims  l.ccii  dIps.mvc.I  to  l.c  icspoiisihit-  i'oi'  ociasionnl 
iKliirnitiltioii  of  somr  of  llu  r/.i.s/nV  loiilaits  illlo  1'  f'SopllilfUls  up  to  the 
level  of  tl.o  licart  or  even  to  tlif  base  of  the  lu'cU.  The  preseiiee  of  the 
L'iistrie  eoiiteiits  in  the  esoi)ha',Mis  starts  a  iieiisfallie  wave,  whieh  pushes 
ihe  material  back  ajjain  into  the  stomaeh.  This  peristaltie  wave  starts 
III  the  ahseiiee  of  any  other  phases  of  the  detrhitition  proeess,  indicatinsr 
that  it  has  lu'eii  excite.l  l.y  the  jyresence  of  the  material  in  the  esophasrus 
Itself,  and  helonfis,  therefore,  to  the  lower  order  of  peristaltie  wave,  as 
seen  in  the  intestines  lint  n.>t  in  the  upper  half  of  the  esoi>hadUS.  Rejjur- 
..'itation  of  food  into  the  esopha<,'ns  oee\irs  only  when  the  intrafjastrie 
pressure  is  fairly  hifih.  It  may  last  for  a  period  of  from  twenty  to  thirty 
iiiimites  after  the  meal  is  taken,  and  disa4>peais  when  the  tonus  of  the 
.phineter  l.eeouu's  iin-reased  as  ;i  result  ..f  the  presenee  in  the  {lastri.- 
iciiitents  of  free  hydrochlorie  aciil. 

y\\H-h  infin'nnition  has  been  seeiire<i     v  listeninir  with  a  stethoseoi>e  to 
ll,r  sounds  ciiiisnl   h<i  sirdiloiriii;!  and   by  obs.'rvinjr   with   the   x-ray   the 
shmlou-s  produeed  alonir  the  eonrse  of  the  esophauns  when  food  impre-r- 
iiiited  with  bismuth  subiiitrate   is  taken.     When   a   solid  bolus  is  swal- 
lowed only  one  sound  is  usually  heard,  but   with   liMuid  food  there  are 
Mvo,  one  iit   the  upper  end.  due  to  the  rush  of  the  fluid  and  air,  and 
the 'other  ni  the  lower  end   (heard  («ver  the  epiu'a^trium'*,   four  or  six 
MTomls  later,  diu^  to  the  arrival  here  of  the  peristaltie  wave  with  the 
iMcompanyint:  oi)eninfr  of  the  eardiae  sphineter  and  the  eseape  of  the 
ilui<l  and  air  into  the  stonnn-h.     Sometimes,  when  the  person  is  in  the 
luni/ontal  position,  this  seeond  soun.l   may  be  broken  up  into  several, 
nidieatinsr  that,  unassisted  by  ijravity.  the  tlni.l  does  not  so  readily  pass 
•liroufrh  the  sphincter.     The  x-r.iy  sha.lows  yield   results  in  eonformity 
ith  Uie  above.     After  swallowini;  milk  a'nl  bismuth,  for  example,  the 
liadow  falls  (iniekly  to  the  lower  end  of  the  esophajjus  ami  then  passes 
•uwlv  into  the  stomaeh.     When  the  passage  of  a  solid  bolus  is  watehed 
Ihe  x-ray  method,  its   rate  of  .l.'seent    will   be   found   to  depeinl  on 
■  'Mther  or  in>t  it  is  well  lubricated  with  saliva:  if  not  so,  it  may  take  as 
as  fifteen  min\ites  to  reach  thi"  st.nnaeh:   if  moist,  but    from  eii^ht 
til  (ijrhteen  see(Uids. 

Vomiting 

Vomitinsr  is  tis\nilly  preceiled  liy  a  feelin-;  of  sickness  or  nausea,  and 
Is  initiated  by  a  very  active  secret  ion  of  saliva.  The  saliva,  mixed  with 
air.  aceumulates  to  a  considerable  extent  at  the  lower  end  of  the  csopha- 
■-'iis,  which   it   disfeinls.     A  forced   inspiration   is  now    made,  duriiif.'  the 

is  iipcii  so  tlnit   th.e  air  eiiti'rs  the  buiL's. 


st 


ns(-  1 


if  which  tln^  ijlattis 


It   later  the   <;lottis  closes  so   that   the   insi>ired   air  is 


sucked   into  tite 


Mi: 


•ii1i~ii 


{    fl 

\ 

3 

!  n 

^^1 

i>  - 

!    i 

-A 

!    I 

;!  1 

•■    !f 

!■  i 

in 

:' 

H 
-  ■  1 

- 

f  1 

. 

t    : 

> 

tji 

i'^ 

'111    r\ 


fit     "i  i     ■  - 


Hi! 


iV'l 


1.1 


1 

;i 

■    i 

4r)() 


i>i(Ji;sTio\ 


I'sopha^iis,  wliicli,  alivjuly  soiiiowliat  ilistciulcl  l.y  saliva,  now  bfcomcN 
markedly  so.  The  alulinniiial  iiiiiscios  then  contract  so  as  to  eoinprcss 
the  stoiiiacli  against  tiic  .liapliiayni  aiul,  siimiltaiiconsly,  the  cardiac 
sidiiiictcr  relaxes,  the  licail  is  held  torward  :iiid  the  contents  ot"  the 
stomach  are  ejected  through  the  jireviously  distended  esophagus.  The 
conii>ression  ol"  the  stomach  hy  the  cont,actin<;  alidominal  muscles  is 
assisted  hy  an  actual  contraction  of  the  stomach  itself,  as  has  heen  elearlv 
demonstrated  l.y  the  x-ray  method.  After  the  contents  of  the  stomach 
its'df  have  heen  I'vacuatcd,  the  iiyloiic  si)hincter  may  also  relax  and 
I)ermit  the  eont(  nts  (  hile,  etc.)  of  the  duodenum  to  he  vomited. 

The  act  of  vomit inu'  is  controlled  hy  a  center  located  in  the  medulla, 
and  tlie  affrnnt  jUxrs  to  this  center  nuiy  come  from  many  dilTerent 
retfious  of  the  hody.  Perhaps  the  most  i)otent  of  them  come  from  the 
sensory  nerve  endin«s  of  the  fa\ie(s  aiid  pharynx.  This  ex])lains  the 
tendency  !o  vomit  when  the  muco:„i  of  this  rcfrion  is  mechanically  .stimu- 
lated. Other  alTereiit  impulses  conm  fiom  the  nnn-osa  of  the  stomach 
itself,  and  these  are  stimulated  hy  inn  lies,  impmtant  amonir  which  are 
stronir  salt  solution,  mustard  water  and  zinc  suliihate.  Certain  other 
emetics,  jjarticularly  tartar  emetic  and  ap(miori)hine,  act  on  the  vomit- 
injr  center  it.self,  and  can  theiefoic  operate  when  ^dven  suhcutaiieou.slv. 
AlTerent  vomitiiii.'  impulses  also  arise  from  the  ahdominal  vi.scera,  thus 
exi)lainiui:  the  \omitin>;  whidi  occurs  in  stian<iulated  hernia,  and  in 
other  ii'ritative  lesions  involviiie;  this  lejiion.  X-ray  ohserx  alions  have 
heen  nia<h'  on  the  movements  of  the  stonuu-h  of  cats  after  the  admin- 
istration of  ai>omorphine  iCannoni.  The  first  cliani;e  ohserved  is  an 
inhihition  of  the  cardiac  end  of  the  stomach,  wliich  ])eeonu's  a  i)erfectl,\ 
tlaccid  hau'.  Ahoni  the  midrejrion  of  the  (Utran.  (U'ejier  contractions  then 
start  uj),  wiiich  sweep  from  tin-  jiyhirus,  each  contraction  stoi)pin<i  as  a 

deep  v'm>:  at  the  hetrinniii};  of  the  vestibule,  while  a  slijrhter  wave  ( - 

liinies.  A  very  stronu'  eouti'action  at  the  incisura  aniriilaris  finalh 
develo|)s  and  comj.letely  ilivides  the  ■•astric  cavity  into  two  parts.  On 
the  left  of  this  constriction  the  stomach  rcnnnns  com|(l<'tely  relaxed,  l)nt 
at  the  ri^rht  of  it  waves  continue  running  over  the  vestihule.  It  is  while 
the  stomach  is  in  this  eon>lition  that  the  sudden  contraction  of  the  dia 
phra'.'in  and  ahdominal  muscles  shoots  the  cardiac  contents  into  the 
relaxed  esoplia>rus.  As  these  .jerky  contra<'tions  are  continued,  tlie  <,'astrie 
walls  s(>em  to  reacipiire  theii'  tone. 


fHAI'TKK  LI  I 
TIIH   MKCIIANIS^IS  OF   DKiKSTK  >\   (('..nt'in 


THE  MOVEMENTS  OF  THE  STOMACH 

The  Character  of  the  Movements 

Hvoii   finiu  the  earliest   <liiys  it    li;is  hcen   recoiriiized  tiuit   tlie  stoiiiaeli 

IHTt'onns     wo   imixd'tant    t'uiictions:    il'    receivinu'   tlie   swallDued   food 

;ind  tlieii  discliaf'riiiR  it  slowly  into  the  intestine,  and  ( 'J  i   initiating  the 

rhemieal  (jrocessos  ot  diurestion.     In  (udei-  to  nndeistand  tlu'  ineclianisin 

liv  which  tlio  stomach  collects  ami  then  disdiarui's  the  I'ood,  it  is  iicccs- 

s.-ii'v  tirst  of  all  to  recall  ceitain   anatciniic    Tads  coin-efninir  the  orjian, 

iind   for  this  jiiirpose   it    is   must    convenient    to   accept    the   descrin^'on 

L'iven  liy  Cannon,  whii'h  is  ilinstiated  in  the  ai'conii)an\  ins:  tifrun       The 

niL'an  is  divided  into  a  cardiac  and  a  pyloric  portion  hy  a  deep  n      a  in 

the  lesser  curvature,  called  the  incisnra  ani.Milaris.     Tlie  cardiac  ])ortioii 

is  I'uither  sulidivided  into  two  liy  tlie  cardiac  orifi<'e.     The  i)art  whieh 

lies,  in  man,  ahove  a  line  drawn  horizontally  thronsh  the  car<lia  is  the 

IiumIus.     The  iiart  lyinir  between  the  t'nndns  and  the  iiu-isiira  annularis 

is  known  as  the  liody  of  the  stomach,  which,  when  full,  has  a  taperin<; 

shape.     The  jiyloi'ic  ixo'tion  l\ini;'  on  the  ii<.dit  oi'  the  incisnra  anffularis 

is  further  divided  into  two  parts:  the  ]>yloric  vestihule  and  the  ))yloric 

'iinal,  the  latter  of  which   lies  next    the   jiyloric  s]ihincter  and   in   juan 

iiHiisures  ahout  '■>  em.  in  hnv'tli   i  see  I'^ij.'.   \'>-i  '  ■ 

The  filled  stonnieh  of  ,i  )>erson  standing  erect  is  so  disposed  that  the 
-I latest  curvature  forms  its  lowi'st  iioint,  wlTu'li  may  he  considerably 
iilow  the  umbilicus.  As  digestion  proceeds  and  the  stomacli  empties, 
the  u'reater  curvature  becomes  irradually  raised,  so  that  ultimately  the 
livhirus  c<pmes  to  be  the  nmst  dependent  part  of  the  stomach.  From 
;ticse  and  many  othei'  observations  it  is  cei'tain  that  the  emptyinir  of  the 
sNiniach  does  not  at  all  depend  on  the  operation  of  the  force  of  irravity. 
Iiiileed,  that  this  can  not  be  the  case  is  iierfectly  clear  when  we  con- 
-idcr  the  disposition  of  the  stomach  in  ipiailrupeds. 

Ilxact  observation  on  the  mo\eMients  wliich  the  stomach  jx'rforms  from 
■'ii'  time  it  is  tilled  with   food   till   it  empiies,   iiave   been   made  by  the 


\  ray  method,  first  introduceil  by  Cannon. 

4.-)l 


Th. 


I'thod 


onsists  in 


f.MMl- 


ii<i 


.Ifl 


Mi 


I 


'I" 


I  i 


)* 


iiii^ 


if\:'' 


,\      ! 


i'^' 


:     H 


i.'t;  , 


|i|(ii:sTiii\ 


inji  llic  .'iiiiiii.'il  with    lood   lluit    Ii;i>  1 ii    iiii|>ici,'H;itc.|   wilh   l.iMiiutli  siili 

iiiti-iitc.  Ilii'ii  cxixisiiii.'  him  tii  the  \-i;iy  iiiid  cither  tiikiiiir  iiistaiitancdiiv 
I)h(>t<>f;riii)lis  of  the  shadows  iir  ohsciv  ill},'  tliciii  liy  means  of  a  (luoiescciit 
sprecii.     Tlie  dcsciiplions  of  tiie  oiiiiinai   h'iscin  ations  niadr  i)V  Caiiiion 


I  IK        1=1 


.I.IM-;     /.■ 


Ill,      -l-HMill,        Al     (       -v     tl„      ,, ■,,.!.:,;     /,     I, III. I, ,^;     I.I.     iihj-u,,-,     all 
;    ''1       l.>l"ii.     ..ir.il;    /',    i,>l.,r  i-        .  |'i,.,ii    Cm 


cm   till'  stiimai-h  of  the  cat   i:a\f  iiccii  so   litlle  modilifd  iiy  olisci'vatioiis 
on  ijian  liial  we  may  tak<'  tlitiii  as  a  coiixciiiciit  t.\  |)c.     In  the  acconipaiix 
illy   li«iirc   I  Kiir.    I'liii    the  outline  of  tlie  shadow    east    hy   the  slomaeh    i- 
sliow  M   at    intervals  of  an   houi-  eaeli   during   cjiufstioii.     Soon   after   tin 


^\^   ' 


FlK-     \^-        lli.iKr,-,:,,-    ..i    ,.„t|„„.    ;„„1     |„,„ti,,n     ,.l     .t„ril.,.  1.    a,    .ik1kj1,,I     I,v     -kiaKtains    takiil    .- 
mail    III    oil'   tri-.  I    ■..>^|M..n     .1    ......,,..,i,    -,.1.  ^      .,    II    .1.        ,1  .."..'*.  ,     . 

A.   iiMiliTa  .ly    1.1    ;    /'.   liia.t,>all>    .i„,,u^      Tli.-   ,l,ar   >,  a..-   at    ih.-   u,.|..-,    vnd   ul    II,,-    sl.imach   is   ,l,i, 

'iir'Tiiri!!!"    "  ''    '""""'   "'•"    ""-   "-I"'""''    hia.l.i.i-    lifs   il„,..    1,1   ,|„.    |„.aii.      ,  |-r,„ii    T     Wir, 


TUi;    MICIIANISMS   Oh"    PKiK.STloN 


4r.:{ 


.t.Miia.'li  liiis  luMM.in,.  iillf.1.  iHTistaltir  wav.s  iiic  seen  t..  take  their 
,„i.'ii,  al.uut  tlir  mi.l.llc  of  tl...  lH..ly  of  the  st-.ma.-h.  an.l  to  .Mmrso 
,„war.ls  the  i.vlo.us.  Ahnvr  tlie  iv-ioM  at  which  tlicse  waves  uncinate- 
,l,at  is,  the  eanliae  half  of  the  l.o.ly  of  the  sto.i.aeh  and  all  of  the 
fwiulus--the.e  arc  iw.  waves,  l.i.l  as  di-estion  proe.e.ls  the  walls  sh.wlv 
;,„d  stea.lilv  eontraet  on  the  mass  of  food.  This  so-ealled  eardiae  poueh 
,lnes  not,  h'owever,  diminish  in  size  so  ia|.idly  as  the  pari  of  the  l.o.ly  of 
the  ston.aeh  over  whi.-h  the  peristaltie  w;i  ,s  are  i.assinji.  The  eireular 
lihers  of  the  walls  of  this  part  of  the  slonuieh  sometimes  ealled  the 
.r,,strie  tnhe  eontraet  tonieaily,  so  that  it  I.eeonies  tnhular  in  f.M'iii, 
with  the  full  .-ardiae  i.oneh  at  the  left  and  al.ov.-  and  the  pylorie  por- 


;i    cM    iMlli    l'""l    mi'i"-' 


;i„n  at  the  ri-.'!.i.  The  latter  portion  meanwhile  .l-es  not  .liminish  miieh 
m  size,  althon^h  the  peristaltic  naves  trav.'lins:  over  it  are  very  pro- 
u.unee.l.     As  will  he  .-h.ar  from  the  ti-ure,  these  ••han-es  in  outline  ^'o 

„  until  the  eardiae  poueh  has  hee.nile  praetieally  empty  and  the  foo<! 
has  heeii  all  moved  alon-  the  now  tulnilar  i.ortion  of  the  body  into  the 
m  lorie  vestibule. 

'i-r«mi   this  deseription    it    is  evi.lent    that    the    funetinn   of   the   eardiae 

.'id  is  to  ser\e  as  a  reservoir  for  the  food,  whieh.  by  a  slow  eontraetioii 

..I    the   walls,    is   jrradnally   .lelivere.l    into   the    fjastrie   tid.e,    where   by 

.'••ristnlsis  it  is  earrie.l  towai'ds  the  pyloric  vestibule. 

The  tinw  re.piired   for  the  i.eristaltie  \vaves  to  travel   from  their  place 

f  nri'„'in  to  the  pylorus  is isid'-rably  lonu'er  than  the  interval  betw.-en 


■Jl 


454 


l>I(il>TI(».\ 


the  uavrs.  so  that  several  ,.f  tluvs.  are  always  svvn  on  the  ston.aeh  ai 
the  same  f.n.e.  Thoy  so.,,  .times  I.eeo.ne  so  protiouneed  in  the  pylori. 
roK.on,  espeeially  in  a  half...m,.ty  stnn.aeh.  that  they  appear  almost  tn 
ol.liternte  the  eavity.     They  always  st„„  at  the  ,.ylorus,  never  «oin,,  on 

to  the  .luo.Ienn.n.     The  rat.,  of  reenrre, of  the  waves  varies  .omewhar 

",'  "■'""'•'■"^   •■'"'"'■•''''■   '"'"!-'  ••'I"""    ^ix    l-r  nite   in   the  eat   an.l   ah„u, 

•luve   ,n  man.     Thei,  initiation  .loes  n„t  seem  to  .lepen.l  on  the  ,.rese, 

"I   .'i-'Ml   n.  th,.  srastrie  .■ont..nts,   for.  when   foo.l  Is   intro.lnee.l   into  the 
stomael,    tlM.y  do  not    wait    for  the   i.'astrie  eontents  to  l.eeomo  neid   in 
'■'■■•"■tl"M    (S,.,.  pau'e  4SL'K      Nevertheless,   aeid   does  seem   so.newhat   to  stin, 
'date  the  d..,,th  and  fr...|neney  of  the  waves,  and  thev  reeur  oftenor  with 
earlH.hyilrate  than  with   fatty  food. 

The    pn.ssnre    in    the   stomael ,tents     //„    intnn,aslnr   ,,rrs,nn     is 

'"^;  ■■'.'"'  •'""^"""  •"  <1 •■"■'li"-  .'nd  and  fanly  hi«h  and  variahle  in  the 

pylone  end  Cn  the  fornn.r  fro„,  (I  to  s  em.  of  water,  and  in  the  latter 
rom  .1  to  :i(),.  Constaney  .,f  pressure  in  the  eardiae  end  i.nlieates 
that  the  stomaeh  wall  must  adapt  itM.lf  very  promptly  to  the  amonnf  of 
food  ,n  the  oriran.  The  hioh..,.  and  nn.re  variahle  pressnre  in  the  pvlorie 
end  ,s.  of  eourse.  .Ine  to  tlie  ,,eristaltie  waves,  and  it  is  intere.stinir  to  note 
that  ,t  .s  snmen.nt  to  prop..]  th,.  -astrie  eont,.nts  thronRU  the  pvloriis  for 
several  e..ntinieters  into  the  duodenum. 


;|;;.1:1!; 


i^'im 


fi  i 


The  Effect  of  the  Stomach  Movements  on  the  Food 

This   h;,s   l,een    studied:    ,1,    hy   .lividin..   the    food    into   portions   that 

;';■"  '''"'■'■'•"•  ■•^-  -"l'"'-l  ""-I.  nl-ter  s, ,!„„..  killin.  ,1,,  ,„i„„|.  tVeeziu.. 

til.,   stoma.-h   an.l   makiufr  s,.eti..ns   .,f   it    (s..e    Fi-.     l.'.T,.    ,•>)    l,,-   ,„.,|. 
mu'  little  p..l|,.,s  of  |,is,nuth  suluntrate  with   star.-h   an.l  .'.l.servin'.^  theiV 
'"■•';'-■"-  un.l..r  the  x-ra.vs:  ..r  ,  :n  hy  renu.vinjr  sampl..s  .d'  the  st"..maeh 
.•"Mtents  l,y  nn.aus  of  a   stoma.-h   tul...   ,R.>hfuss  tul.e)    inserted  .so  that 
.ts   fr,.e  eu.l    In.s   in   eith..r   th,.   eanliae   or   the   pyloHe    reudon.      Hv    th, 

hrst  of  the  ahove  m..thods  it   has  I n    foun.l   that  the   first  monihft.ls 

nt  f.M..l  he  ahuii.  the  i:reat..r  ..urvature.  wh..re  th..y  f.,rm  a  Inver  ov,., 
whwh  that  sul.s,.,|u,.nlly  swallow,.,!  a,.,.uu,ulat..s.  with  the  last  ,.or- 
ti-ns  n..M  th,.  ,.ar,li;,.  Tl„.  p,.psin  an,l  hy,lr,...hl..rie  a-i.l  ,d  the  .-ar 
•li-i..   en.l.   ther..|or,.,   aet    soon,.st   on    the   first    swall..w...l    porthu,    ..f   . 

!""•'''•  '"""^  ''"■  """•'■  •■' "•'.^-  ^"allow..d  ....ntral  mass,.s  are  not  afT....t..d 

I..V  the  seer..f,ons  for  son>e  time,  so  that  opportunitv  is  iriven  for  th.. 
saliva  mix,Nl  with  the  f.,.,d  t..  dev..l..p  its  di-.'stive  aeti.m. 

As  has  heen  sh..wn  l,y  r-niovinir  th,'  sfoma.'h  eontents  with  a  ttd)e  at 

varu.Hs  peno,ls  af!,.r  f.-edin^  v.Uh  .Un-rhv  f 1.  ..,M,si.l,.rai.ie  amvl.dvsi. 

.nay  oeeur  for  some  ti,u...  When  separate  .sampl.-s  are  remove.Mn  this 
way  from   th..  eardiae  an,]   pyl,u.ie  parts,  it  Ims  l...,.n  found  that  after 


f® 


Tin;  Mi;<iiA\isMs  oy  i.itiKSTHiN 


4r)r) 


half  an  lu.ur  the  contents  of  l.oth  havo  about  the  same  iHTcentaKc  of 
su-ar,  l.ut  that  for  some  time  after  this  interval  the  cardiac  contents 
contain  consi.lerably  more  sujiar  than  the  pyloric.  Later  the  percentages 
„f  suu'ar  aiiain  become  about  cpial,  no  <lonbt  ,m  account  "f  difTusu.ii. 
The  (liastatic  action  in  the  fundus  is  (inally  brou<;ht  to  an  end  when 
,he  contents  become  completely  periM.Mled  by  the  hydrochloric  acid. 
I„  tills  connection  it  is  worthy  of  note  that  the  addition  of  hydrochloric 
arid  up  to  the  point  of  neutrality  «rcatly  accelerates  the  rate  of  diastatic 
digestion. 

As  the  otiter  layers  of  f 1  in  the  stomach  become  partly  disjested  on 

account  of  ihc  action  of  the  pei.sin  and  hy.lrochloric  acid,  the  food  is 
slowlv  pressed  ini,.  th.'  active  ri-h*  half  of  the  stomach,  nhere  by  the 
action  of  the  peristaltic  waves  it  is  movc.l  on  to  the  i-yloric  vcstdmlc. 
Hy  observin-  the  x-ray  shadous  east  by  two  pellets  of  bismuth  subni- 
tratr  it  has  been  noted  by  CannoM  that,  as  the  peristaltic  wave  approaches 


1  ii!.    15".--Sviti"li 


,,f   the    l-.j/.M   >t.'iii^iih    ir:itl    S'.im-   linu-    altrr    f.  .-ilini;    "Uli    '• 
ilirttrc  Illy    c.lo-.v.l    ].'.lli'iin.      (Frum    llrwill's    l'l!\  siuli'ii:  .) 


■A    RiM-n 


three 


a  pellet,  it  causes  it  to  move  forward  more  rapidly  for  a  short  distance, 
l.ut  soon  overtakes  it  and  in  doinsr  so  causes  the  pellet  to  move  back  a 
little  towards  the  fundus.  This  backward  movement  is  less  than  the 
forward  movement,  s.,  that  after  the  wave  has  passed,  the  position  of 
ihe  ,.ellet  is  a  little  forward  of  that  which  it  woul.l  have  occtipied  had 
-here  been  no  wave.  The  b.havi.U'  of  the  pellet,  ami,  therefore,  of  the 
-lomach  contents,  is  very  like  that  of  a  cork  tloatiufi  at  the  ed>:e  of  the 
-a;  as  each  wave  approaehes,  it  hurries  the  cork  on  a  little,  but  after 
"s  passajrc  the  c.u-k  r les  airain  until  the  s« ud  wave  carries  it  still 

1  little  farther  forwanl.  As  the  peristalti.'  wave  ai.proaches  the  pyloric 
vfslibule  and  beconu's  more  jiowerful  its  ctTect  on  the  pellets  becomes 
more  marke.l.     Thev  are  carried  rai)idly  alonu'  *his  part  of  the  s..,mach, 

mtil  the  pylorus  is  reache.l.     If  this  remains  closed,  they  are  shot  back 

iito  the  vestibule.  From  nine  to  iv\elv,-  minutes  ma.\  elapse  before  they 
,i.v  transferred  to  the  pylorus  from  the  place  where  they  are  tirst  affected 

!  y  tlip  peristaltic  wave. 


I  ii 


mii 

ui 

IB' 

|U| 

H 

;» 

r- 

jB 

B 

i 

i 

1 

y  ■ 

ii 

h 

M 

rli 

}Wr^ 

1 

1  1 

:\    ■ 

|H 

f 

, 


ii^' 


456 


l>i(fi:sTit)\ 


Thcs..    ol,M.rMi1i..iiv    ina.l ,    cats   iiri.l    (.thcr    hilmratoiv   aniiiials    ii. 

'I"ul>»  also  api.l.v  in  flic  rfiM..  of  m„im.  rinn..vnl  of  tho  <-oiil,.Mts  ..f  tin 
•■"•i''«lif><'  iui.l  i.vloii,.  n.(ri..Ms  scpanitrly  with  a  stoii,,,,-!,  tul.c  i.ftrr  fcxiin- 
"itli  ;i  t.'st   moal   part   nf  ,vhi,.li   was  colonMl  wifl,  ..arinin,.  nr  Hiarcoal 

•'f  '*''"""  "'"'  ' '  "•'  "'<■  ••"loiiiitf  inatcfial  was  present  in  tlu nt(>nt> 

"I  tlie  pylori,,  en.l  up  to  tw..ntv  Miiniites  or  s(.  after  the  foo.l  l,n,l  l.een 
ffiken.  Ft  then  appeare,!  In.t  at  lirst  only  in  traees,  A.n.tl.er  imi.ortaiil 
.l.stinetion  l.etween  tl,e  foo.j  in  the  two  portions  of  the  stomaeh  relates 
I..  Its  eonsisteney.  In  the  pylorie  en.l  it  is  semifluid  an.l  homoRen..OMs 
.'"  '•'•"'■"•''•'■:  I"  flu.  eanliae  en.l.  on  the  otli.T  han.I,  it  is  a  lun.pv.  rathe, 
incoherent   mass. 

The   -astri,-  inoven ts   must    irreatly    f.irilllal,    tin    ,i;,,<si;rr   ,,n,rr,u. 

'"  *'"■  '' ••"•''      '"   ""•  '•■"•'li='<'  l'i"-t   the  nn.listurl.e.I  condition   of  the 

t""d  will,  as  w,.  liave  se..n,  faeiiitafe  the  di-estive  aefio,,  of  ptvalin 
"hneas  ,n  the  l.o.ly  of  the  st.nr.aeh  the  peristaltic  vaves,  l.esides\noN . 
iML'  th,.  food  onward,  will  tend  to  hrin^r  fresh  portions  of  nnieous  men, 

'";■""■■""'   '■'""'   '"'•'■  -"  ♦'"•<   the  latter  hee.nues   more  thoron-diK 

.M.xed  w.th  the  ,.epsi„  and  hydrochloric  aeid.  In  the  pvloric  part  wlicv 
>M.  hy,l,„ehlor,c  a,.id  is  secvted.  the  contents,  alr-adv  suffieientlv  aeid 
m  reaction,  hcome  nioiv  thoroughly  churned  up  with  the  local  pepsi,, 
se..r,.t.on.  so  that   proteolytic  acti.n,  i.rofrresses  very  rapidly. 

The  peristalti..  M  avcs  aNo  farililolf  ahsor,,l!on  fro,,,  ll.r  sh„„„rh  of  sud, 

siil-stanees  as  uducose  i neenf rated  solution  ai„l,  pn.l.ahlv    „f  hvdn, 

Iv/.'d  protein;  wate,-.  however,  is  not  al-.sorhed.  One  efTect  of'sud, 
=rl.s,n.ption  is  the  ,.roduction  of  ,.aslrin,  Mhieh  we  have  seen   is  the  hor 

;;'"""  ':;"■*•;.■ '  ;"    •""'"•"i-iMU-  the   s:ast,.ic   secretion    after  the   psvch;.- 

II-w  Th..  tan  that  the  mu....sa  of  the  v,.stil.ule  has.  relativolv  t.",  tl„ 
-•.•inliae  en.l.  f..w  s..cr..ti„j:  .dan.ls  is  in  ha,-mony  with  the  view  that 
.•d'so,pt,on  ,s  a„  in,po,tant  function  .,f  this  i.a,t  of  the  st.miaeh. 

THE  EMPTYING  OP  THE  STOMACH 

The  Control  of  the  Pyloric  Sphincter 

When  .iis:..stion  has  pro. Id  far  en-n.^h  in  th..  st..maeh  t.,  hrin^r  the 

' '  '■""  "  ''■"""-"■"••""^-  -"'I'lik.'  Ilui.l   .chyme),  portions  ..f  this,  as  ll,.  . 

iiiv  .Irivcn  a,^,n,st  th..  pyloric  s,d,in,.ter  -,y  th."  p..ristaltic  wav,..s.  instead  f 
iM-injr  return..d  as  an  axial  str..am  into  the  st..n,aeh.  are  e.i,...te.l  int..  th.. 
.ni.idenuni. 

We  n.u.t   now  consider  the  meehanism  ),y  whi.-h  the  pvLu'lc  sphineter 
<'l'.'Ms  t,.   permit   th..   pa.ssa.rc  of  th,.  ..hym...     Il-n,l,ar.im,.„t    l.v   the   peri 
stnlt.e  M-aves  is  ,.vi.I,.n,|y  ,„„   , aiise  of  its  .,penii,,r.    f.u'.  as  ^^  e  hav.. 


Tin;  .Mi:i  HAN  ISMS  ok  KiiiisrioN 


457 


MIL- 


Ilf> 


ml 


ills 


*• 


I  (11,  iiiiiny  s 
\  iilcullv 


url 


1    WHVt'S    llli'\ 


iiri'iM'  fi 


iriliT  ttiiil    tlic   iiitcstiiif   may 


liclmcd  witli  liiiiif  musses  n 


t  it  witliout  tills  result,    Siiiec  it  is 

not   smMeiily   Ik me   "vor- 

•iisioiially 


r   I'.kmI  tliiit    tlie   pyliHils  only 


it    mitrlil    lif   tlmiiulit   tlmt    its  niiei 


It:  (lepeiiils  upon   the  <liston- 

It  is  tnio  tliat  t«xfes«ive  distcii- 

f  tl..'  uppc'  imrt  of  the  inte>tiiie  .lo-s  l.oM  the  pyloric  spliinetor 


.i|iens, 

lion  of  tlu'  upper  ) 


lioii  o 


,(  tlie  intestine, 


sell,  but  tliis  ean  im 

mtities  ol'  eliyine  are  never  I'liund  here 


1.  lie  the  physic'lo'_'ie  stimulus,  lieeause  ( sh 


leral.h 


The   first    elue   to  tlie   re 


il   nature  of  the  meehaniMii   "as 


ifl'oiileil   by 


iliservinff  the 


(■havior  of  the  sphineter   win 


solutions  art"  mtn 


iluei'il 


ito   the   iluoilennin 


throuu'h    a    fistula. 


alise  a  eonip 


lete  inhibition  of  ".'astrie  eva 


Aeiil   solutions   were    found    1<> 
•nation,  whereas  alkaline  solu- 


Mnns 


hai!  1 


lo    (' 


ffeet.     This 


(lifTerel 


lei'  indieates  that  aeiils  in  eontaet   wi 


th 


'lie  duo<lenal  inu<' 


membrano  relle\ 


|\  exeite  eontraet .on  of  the  sp 


hinc- 


ti-r,   aiK 


I    that    it    relaxi's    cnily    after    the 


d     has    beeoil 


le    nel 


itrali/i'<l 


i\   ni 


ixini:  with  the  panereatie  .jun 


e  and  bib' 


On  a nnt  of  the  uieat  iniportanee 


if  the  pyloric  mechanism  in  insiir- 


that  the  ch\nn'  shall  enter  the  intestine  only 


sucli  c(uantitu's 


that 


it  <-.in  be  inojK'rlv  actecl  upon 
,!!■  of  interest  to  consider  brii 
liv  which  this  nn'idninism  has  Ir 


bv  the  intestinal  dit'cstiuir  .juices,  it  aviI 


fly  some  of  the  exi)erim 

studieil.     We  may  eo 


ental  observations 


isider  first   the 
(I  rrhi.nttuiii 


.idence  that  '/././  on  Ih,   slonnirh  si,h    of  tin    piil^inix  cinrvv 
.,'  ih,-  s,,hn,r',r:  I  li   When  earbohy.lrate  food  is  fed,  it  ordinarily  leaves 
the  stoma.di   t;-Mrly   rapidly,  but   if  its  acid-al.s.nbin-  power  is  ircrfase 


iv  mixmp  i<   wi 
lelaved.     (2')    Proteins 


th  sodium  bicarbonate,  exit   from  the  sloma-h  is  Krea 


itlv 


ordinarilv  leave  the  stomach   more  slowly  than 


■nrlio 


livd rates,  bii 


but    if 


d    jiroti 


are    fed,   their   exit    is   much   1I 


lore 


ipid,     CO   If  a  fistula  is  made  into  the  p 


>vloi'ic  ves 


)f  the  contents  can  be  removed,  it   wil 


tibule  throuirh  \\hi<di 
found  that  .just  prior  to 


lie  opening 


if  the  pyloric  sidiincter,  a  distinctly  aci 


1  react i'lii  develojis 


the  food;  and  furthermore  it  acm  so 


liition 


injected  lhrou','h  this 


ula.  thev  cause  the  pyloric  sphincter  to  open,  whereas  a 


lUali 


es  re 


liufl.     (4^   A  similar  ( 


Its  oppl 

II'  demonstrated  by   applyi 


tard 
ffect  of  acid  in  op''nin<i  the  sphineter  cai 
it    to   the    jiylor 


nicosa   oi    an   exe 


ispd 


laeh  kejit  alive  in 


oxvjrenated  b'imrer's  solutio 


evK 


leiice 


■I'he 


ihinrtrr  is  as 


that    arlil    on    IIk    (liiodrmil    siih     innxix    closin-r    of    ihr 
follows:     I  1  >  When  acid  is  placed  in  the  duodenum  tlirmi<:li 
11 


fistula,  the  sphincter  will   not  o 


ipcn:   (2)   when  the  pancreatic  ai 


id  b 


cts  are  lijjated,  the  stomach  emi)ties  m 
d  f:n  the  discharge  of  i)roteMi  is  consu 


icli  more  slowly  than  normally: 
lerablv  hastened  if  llu-  i>ylonis 


sutured  to  the  it 


on  It  was  o 


testiuf  below  the  <luodeniim.     After  smdi  an  opera- 


bserved  that  the  protein  be-^'an  to  leave 


the  stomach  throll^dl 


i 


i   t 


ii; 


Vm  'M 


1  ii. 


I  i.f 


iii< 


458 


DKIKSTIOX 


tho  i>ylori(-  sidiiiicttT  iilx.iit  the  satne  time  iis  iioniifilly,  Imt  tlip  su1)M' 
<iueiit  fvaciiatinii  was  vcrv  iiuk-Ii  awchM'afi'd,  Ix'cnvisp  no  acid  came  n, 
{•iiiifact  witli  the  (lunilonal  niticdsa.  Wati-r  aixl  vuk  "liiti-  may  U'-.im 
tlip  stimiai'li  iiMlfiiciidciitly  of  any  acid  rctlcx  coiifftil  of  tiip  |)yl(iriis.  |'.\ 
oliscrvations  made  tlin>iii.'li  a  diiiidi'iiai  fistula  it  lias  iiccii  found  that. 
after  a  <|uantity  of  water  lias  lieeii  swallowetj,  most  if  not  all  of  it  ver\ 
soon  enters  the  duodenum  in  a  more  or  less  continuous  stream.  It  is  ii<> 
douht  on  this  account  that  driidunn  contaminated  water  is  especialh 
danifcrotis  on  an  empty  stomach. 

The  iiirroiis  ixillniiiii  lliroutrh  which  these  acid  rellexes  take  jdacp  lia> 
lieen  shown  to  he  the  myenteric  ple.\ns.  Indeed,  the  wliole  nu'chanism 
is  (piite  analoirous  with  that  which  we  shall  see  occurs  in  the  intestin. 
diirini:  peristalsis:  the  stimulus,  that  is,  the  acid,  causes  ii  contraction 
of  the  };a.stric  tulie  lieliind  it  and  a  dilatation  in  front. 


-IK.  l~^.  (),,il,,„s  ,,f  ,|,a,l,)w.  III  ai..l..„an  ,.t,l,imc.;  I.y  ,x|„,-iiri.  I.,  xrays  J  li.nirs  .ill, 
IfclnlK  Mill.  l.""l  .-.iluining  iM.im.lh  siiLtiitrau-.  Tlu-  f.,.„l  in  A  ».„  Iran  l..-cf,  and  In  /(  linil, 
ri.e  Ih.-  .ni.il.T  „/.■  ,,t  ilu-  M,M,ia,h  sha.L.w  an.l  llu-  iniuh  Kr.atrr  total  arra  „f  III,  inU-sUu 
>ha.|o«s   II,    /(   ,!,.,„    ,„   .1    »i,.,„.   ,|,a,   c.irl.,.hy,|ratf   Uav.s  tli,-   •.(..mach    .arll.r    lliaii    |.rot<-i.i. 


I  Fr,. 


Rate  of  Emptying  of  Stomach 

The  relationship  of  those  facts  to  tlir  rati  at  which  fliflnrnf  foodstiiils 
harr  thr  stomach  is  very  readily  explained.  The  method  for  invpsti-at- 
mij  this  prohlem.  which  asraiii  wo  owe  to  ("aunoii,  consists  in  feediiif;  ani- 
mals with  a  strictly  uniform  amount  of  diflVrenf  foods  made  up,  a- 
ne.iily  as  i)ossilile,  of  e(pial  consistency  and  containiii!,'  bismuth  suhni 
trafe  in  the  proportion  of  ;")  f;,,,.  to  paeli  2r>  o.c.  14y  feodinjj  such  mix 
tiires  to  cats  previously  starved  for  twenty-four  hours,  and  examiniuL' 
the  alMh.men  liy  the  x-ray  at  re-:ular  intervals,  the  shadows  ca.st  l>y  the  food 
after  passage  into  the  intestine  can  he  outlined  on  traeiiis;  paper,  and 
the  total  length*  measured  Hmj.'.  in.Sl  Tn  takinj;  this  as  an  cstiinatc  <'• 
thp  amount  of  food  in  the  intestine,  several  errors  are  no  doubt  ineurred 

■This  IS   pfrm!»si!,lr  -m.  c  th-  s|,a.i..«>.  an-  |.ratlKal!y   -II   „f  the   sami-   wi.ith. 


1111.    Mil  11AMSM>   <'!■    I'liil.MloN 


4r)9 


„„  account  of  the  ci-oHsiiiK  im.l  foic>li(.it.-iiiiii.'  '>f  thf  loops,  .•tc.  I.ul,  as 
llu-ir  coi.staii.'v  tr^titics,  Wwvv  in  no  .l-ul.t  tliat  tlio  results  arc  sumciontly 
.■lose  for  the  purpose  of  fiiuliiiir  out  h(.\v  (|uiekl.v  foo.l  Kiiins  n.-ess  to  the 
Miinll  intestine;  and  the  method  has  a  Kirat  advantage  over  all  others 
,M  that  diirestion  is  M,>^^v^  to  W'^'rM  pra-tieally  without  interruption, 
rh,.  points  we  have  to  .letermine  are:  (I)  when  the  food  first  leaves  the 
^toInneh;  CD  the  rate  at  whieh  dilTerent  foods  are  disehar-e.l;  CI)  the 
time  re.|uired  for  the  passaije  through  the  small  intestine. 

Let  us  eonsi.ler  first  of  all  the  results  ol.tain.'d  /-/  /.../„.-,  ,v>th  prw 
t„alhl  l»in  fat  or  ,arholn,,irntr  or  ,,roln„.  H.v  |.lottin..'  the  leM-.Mh  ot 
the  shadows  in  eeiitimeters  alon-  the  .ndii.ates.  with  hours  alon«  the 
ahseissa-.  curves  such  as  those  shown  in  l-'iir.  m  have  l.een  M..Mired. 
When  fats  were  fed  (.lash  line  in  ehartV  the  disehar-e  l.-nan  rather 
.Inwlv,  an.l  eontin.ied  at  a  slow  rate.  Kven  affr  seven  hours  some  tat 
still  remnine.l  in  the  stomaeh,  and  at   no  time  was  any   lar-e  Muantity 


i 


,:,„.     ,V,.-a,rve<    ,o    show    U,..    ---",  ='«''-«^"';,,''-l|;  .^MJ-.J^^^/ir  ^., 'l^    l^i.^'i^" 
.lash    lini-.    for    prntrin    fo,..ls    in    the    h.avy    h.i.-.    .i.-l     l-r    i  .irl...lu.M.i'. 
I  l-rnm    I'anjiiin. ) 

present  in  the  intestine,  indieatins;  that   almost   as  (luieUly  as  it    is  dis- 

rharced  into  this  part  of  the  j;astrointestinal  traet  fat  l.ee.m.es  difjeste-l 

and  ahsorhed.    The  diseharjie  of  ..arl.ohylrates  was  .,uite  difTerent  (  n.'ht 

line  in  chart  1  ;  it  !•    ran  often  in  ten  minutes,  and  so,.n  l.ecame  almndant, 

n.achiuK  a  maximum,  as  a  rule,  at  the  end  of  two  hours,  after  which  it 

Mil  off,  the  st.nnach  l.eini,'  em|.ty  in  ahout  three  hours,     l-rotem  Ictl  at  a 

rate   interme.liate   between    that    for    fats    and    that    for   carl.ohy.lrates 

,  heavy   line^.     I/ittle   left    l.efore    the   first    half   hour;   the    curve   then 

slowlv  rose,  attainiiis:  a  maximum  in  alxmt  four  hcnirs,  and  then  -radu- 

allv  declining  at  about  the  same  rate  as  it  rose.     It  is  interesting  to  note 

that  at  the  end  of  half  an  hour  about  eii-dit  times  as  much  oarbohydrate 

i.iid  left  the  stomach  as  prohi,,;  at  tlu  end  of  an  hour,  live  tin>es  as  much. 

These  results  are  elcarlv  depend.nt  upon  the  rates  at  which  the  dit- 

icrent  foodsvuffs  assume  an  acid  reaction  in  the  stomach.    Carbohydrate 


tk 


4tW 


l>l<il  .^I'ldS 


liiiN   Mil   i-i>iiil>iiilliu    |M.\(,i-    l'..i'   a.'iils.   so    tiiat    Ihf   iiriil    M'.Trlf.l    ujtli    ll,. 

Jisychic  jiiiiT  iciiinins  i iiiiMiircl  nnil  on  lijiiniiiir  tin-  pyloric  Vf«itilpiil. 

'•''•''••'^  •' iH'iiiiiu  1.1'  111.'  spliinrt..|-.     I'loiriii,  on  the  oih.r  Iwind,  n-  i- 

well   knouM,   ;ili«.orl(s  consiilcinlilf   i|Miinlitii's   of   I'ln.    Iiydnicliloric   n<'i<l 

so  fhnl  forjionu'  .■oM>ii|<Tiil.l.'  time  jillci-  it  is  tiikcn,  I'  llic  nriil  oxisis 

ill  a  frci-  Ntiilr.     TiiIm  owo  ihcii-  slou    ilisi'li,ni.'<'  paitly   to  inliiliition  oi 

jriisfric  MTiTlion.  iincj  pjirtly  to  ilir  lon-.'ri'  tiinr  it  liikcs  for  tlinn  to  Ik i, 

iii'mriili;'!'.!  in  llir  iIimhIcmiiiii.  I.c-Mnsc  of  tlu^   f.ittv    iiciil  split  off  hv  tin 

IM'tioM   of  lipiisc, 

Flltrrestim;  oliscr\  ;it  ions  luivc  also  Krcii   niioli'  on   ti,        .ttr  of  discliaiL"- 
«lifii   \arioiis  ,„ii,l,ii„ili„„i  „f  ff„„lsli,ils  w.Ti-   f.'il.     This  lias  liccii  ilon< 

'•>    ''<<"liiii-'  I" IntiilT  l.rfor.'  til.'  oth.T,  or  In    niixinir  tli.'  f IstiilT. 

Wlicii  i'arl.o|i>.iral.s  wcrr  fi^.l  first  an.l  tlicn  prolr'n,  tli."  <lisc|iar(;i.  \>r 
jraii  ninrli  earlier  tliaii  witli  protein  alone,  I.eeaiise  tli.'  earlioliydrale  f i 

lirst  r.'a.-|ied  flw  p>lol'ie  \..uil,uie  ise..  paue  4:.4  .  l|o«e\,.r.  at  the  I  II. I 
of  twc  hours,  when  th.'  earhohydrate  eiir\e  should  |,e-;ili  to  eoine  dou  n, 
it  remained  hiirh,  indiealiii!.'  that  the  protein  had  l.y  this  time  iciielie.l 
fho  pyl.iiis  and  was  heiiii:  diseharsjed  at  its  own  rate.  When  the  meal 
was  fed  Im  fore  the  eiirh..liydrate,  the  eiirve  to  start  with  was  Pvaeth 
like  that  (>>]■  protein,  lieeomiir  however,  i-oiisiderahly  heichti'iied  later 
when  the  earhohydrate  reaidie,.  the  pyloric  vestihiile,  Tlie  preseiiee  oi 
protein  near  tli.'  pylorus,  therefore,  distiiietl\  retards  the  evaeiiatioii  oi 
earholivdrate  from  the  stoma<'h.  These  facts,  it  will  he  remarked,  al! 
lit   in  adinl.al.ly   with   the  ohservatioiis  which   n  ••  in:    ,    already  dr'aile.! 

c-oneeriiiim-  thi'  disposili f  food  in  the  stomaeli. 

^^''"' iMiires  of  eipial  jiarts  of  difl'erent    f Is  weiv   fed,  the  resiilt- 

indieated    that    the   emptyiiii;  of  the   stomach    -urred    at    a    rate    which 

was  intermediate  hetween   those  of  the   foods  taken   separately.      Mixin- 

protein   with   eari.ohydriite,   f(n'  example,  ; leiated   the   rate  at    whici 

protein  left,  and  inixiMi;  fats  with  protein  caused  the  protein  to  leav. 
the  stomach  ciisiderahly  imn-e  sloul\  tiiaii  if  protein  aloiu'  liad 
heeii   feci. 


m 


I- 1    I 


Influence  of  Pathologic  Conditions  on  the  Emptying 

An  impinlant  siiri:ieal  appli,',ition  of  these  facts  leeriis  llu    h,h„r>,., 

<,f  fooij  „fl,r  u'lslnxnln-oslnnHf.  It  has  heen  tliouirlit  that  this  operation 
would  cause  the  food  to  l.c  drained  fmm  the  stomaeli  iiit<.  the  intestiic 
and  thus  leave  the  reyioii  of  the  stomach  l.etwceii  the  listiila  and  tin 
I<yIorus  inactive.  This  assumption  is  l.ased  on  the  id.'a,  whieli  wo  ha\. 
seen  to  he  en-..ne„us,  that  -ra-.  ifv  a.ssi.>ts  ii,  ll,,-  emplNin-  of  the  stomaeli 
As  a  matter  of  fact,  it  has  heen  found  that,  if  the  -rastroentero.stoniy  i- 
ma.Ie  when  there  is  no  oLslrnetion   .-it    the  p.\  h.i  us,  the  chyme  takes  it- 


..     Ui 


Till 


Ml  1  II  Wl^M-   I'K    "li'l  -ntis 


1*11 


\M 


,,„„„|    |,Mss„u.'   lln..uL.|,    llM.    >,.l.iM-..r   .imI.   nl.H..si    uill t    rvrpt  mii. 

|,,,v.s  l.N    \hv   INt.ihi.      Wlin,   111.-   |.>l..n.H   i.   imillx    lu-l.-l.   Hm- 

, 1  soiiiHinirs  ,msH..s  in  ll.r  ..-..I  nv.'.v.  ""'I  v.H.rti.n.s  l.v  tl,..  .Inn.a.Oi. 

,• ,,„«■  r..r  ll.iH  ,„v.lilr.-r,..M   f..r  tl,..  |.>l..n.'  patliun.v    .l.p.Mi.U  ..,,  tli. 

,„,„,„,.,•   rnn.litioi.^   ill    tl..'   UmM'm-   ,-..Ht.iits.      .i„,tru,.nlri-..HtH„,y.   Ilirn- 
,„,,.     is   rtli.-inil    oiiK    «Imm    UIhss    Mir,-l,i.niri.l    ..l.>l  nirt  Inn    rx.sts    at    Hh' 

pvlonis      Til.'   ..,Hiatio„   sl,..„l.l    ii.v.T   1...    ,..Tln,,nr.l    in    tli-'   Ml.sri, t 

,|,.|iic.ii>triil.l<'   .>rL'iiiii«-    I'viniic   cliMiis.' 

\„.,tlirr  „l.i..Hi..M   In   unstnHlil.l  n>tnn.  V    in    tlir   l.lvs.n.T   nl    „    iiMlulou-. 

pvlnii.  spliinrl.T  .-.■sKnii  Hi.-  faH  tl,al  Hi-  C I.  altrr  |.a-,M«  Hir  M.l.m... 

„,,  .,,,.1  .....viiiK  alnHU'  till.  iiilrstiM...  li.av  at-aiii  '•ntrr  tlir  slmnarli  tlin.null 

,1,..  fistula      Tl.is  is  iiinst    liU.'ly  t ni-  nv1i-m  Hi.    sfina-'l.   w   lull   "I 

,• 1    )•„,•  uimI.t  tlirs.'  .•nn.liti.Mis  ill.'  st .•.■l.-liin--'  «(  its  nails  s,.|,arafs  Hi.- 

,,|,,,s  ..!•  II l..'.iiiiir.  II..'  iiil-'sliii.'  l..'ii"-'  '!-■''""  """  ""•'"'■'"  " 't";-- 

.„  that  111.'  npniiii..'  li-t".-i  tl«''  -'""'^'-'l'  =""'  "'"'  '""•'*"'"•  ='-">ii..s  111.- 

,„,.„,  „f  lu ur.iu    slits,  uir..-li   a,M    liU.'  vaK.-s  p-niiitli,,..'  tl...   It" 

,,„..,•  hut    p..-vn,li..«  its  .-ap.'   irn.i.  tl..-  si arh.     Cilv   s.. i.ln.n   uii.I.t 

,i„.s,-   ..im.n.sta.i.M-s   .'an    aiiv    fn...l    |.ass   int..    tlu-    luLstii,,.    h...v.„i.l    th- 

MoMIM.-h     np..ui,...'.       K-p.-at-d     vnlMiUlM..     a.t.T     UMst  rnrlll -mstnlMV      has     h.rll 

„|,s..rv..,l  ill  .•sp.TiiM.i.tal  animals  nni>  xvli.u  nh«tni.-t,M-  kinks  nr  nlh.-i 
,l,,„.,nslrahl.-  nhsla.-l.s  u.-r-  pi.-nt  in  Ih.-  .„'Ut,  th-  nl.stni.-l.nn  h.-mi.'  In 
,.^,„.,,  i„  „„.,  ,„,,  „,•  ,1,.,  infsiin..  iM'xnn.l  lis  Mlta.-hiii.-nt  tn  tin-  slnuia.  n. 

Wh.-n  III.-  pvl.ni.-  ..hstiu.'ti.M,  is  ..nnipl.t.-.  r..n.l  must,  ..f  .■nurs.-.  U-av.- 
l,v  ilw  tistula.  .li.^M.sljnn  hv  th.-  pan-irali.-  .juic..  a.i.l  hih-  h.'.nu'  st.ll  .-ai- 
,;,.,   ..„   !,,,-a.is,-  nf  tl...   la.-l    ;hat    I-'  a   .•niishl.-iahl.-  .lislaii.v   .h.wil   Ih.' 

.,„.-slii,.-.    s....n.|i..,    uhi..h    u..    I,a^.     s.-, ss.-ntia!    Inr    th.-    M...n...iM 

,„•  ,l,..s..  Ilui.ls.  is  slill  ,.in,lu.-..l  hv  tin-  ......a.-t  nf  ,h..  a.-i.l  .•!.>»..-  uilh 

I,.,    int.-stinal    ii.u.-nsa.      ru.lh-r    pi-vi^ion    for    aih-M-iat.-    .liu-.-stinu    nl 

: I   in   su-h   .-as,,   is   s....u.v,l.   as   sn,...-   of   tl..-    f 1    aft.r   l.-avu..„'   th- 

■i.tula  l.ass..s  ha.-k  f.-r  snuu-  .listaii.-.-  iiHn  th.-  .Iun.l..,...i.,  "h.-iv.  hnw.'V.-.'. 

.    s. .-x.-it.-s   p.-,istaUi.-   uav.-s.   uhi.-h    a-j^-ii.   .-arfv    it    fmuanl.       Ihis 

.,sun-s  th.M-n.mh  n.isiim  uith   tl,.-  .li-.-.-sliv..  .jui.-.-s.      I'.-nm  th.-.r  .-xp...- 
,„.,„„1    ..xp.-ri.-n.-.-    fai.iu...    =n„l    lUak.-'      .■.-nmMi.-n.l    that,    ^vl..-u    t  i.- 

-Mula  has  In  h.-  ma.!.-,  it   sh-mM   l-  '«-  laiL'.-  as  pnssihl."   an.l   n.-ar  Ih.' 
l.h.n.s,    aii.l    that    th.-   slo.na.-h    aft.iu  ar.ls    shnul.l    n..t    he    alhnv.-.l    U> 

,,;.„„„„,  tili.-.l  uilh   f.....l.     T..  avni.l  kinkinu^  nt  th.'  '-^,1.  th.-y  alsn  .-.-.-..m- 

„„,n,l  that  s,.v.-.al  .•.-.itiiii.-l.-is  nf  th.-  ii.l.slin.-  sh,.ul.l  h.-  alta.-h.-.i  tn  th.- 

^tnn.a.-h  .listal  tn  tin-  ai.ast..iiinsis. 

Tin-    .-flV.-t    of    huixrnri.hlu    nf    tl nul.-.its    n.l    til.-    .-v-Vt;  in-    n''    thl" 

,,.„„a.-h  has  h.-.'n  stu.li.-.l   h>    f.'.-lin--'  animals  wilh   p.Ha.n.-s  .-..i      ■    Ui'A 

^al■vil|.'  p.-n-.Mila.,'.-s  ..f  hy.ln..-hln.i.-  a.-i.l.     With  an  a.-..litv  ..t   tV-.  V^'V 


t 


I 


462 


MKil.STIO.V 


fll:;,  .  ' 


M 


!*• 


ciMit,  the  liitc  ol' ilisclijirp'  was  iiicifiiscd,  lint  it  lici-aiiic  slower  wlirii  tin' 
Jii'idity  rosi'  to  I  per  cent.  With  an  acidity  of  0.5  per  cent,  the  rate  m 
discliar^'e  was  aliont  tlie  normal.  Hyperacidity,  therefore,  causes  a  retar 
fiation  of  the  tinptyinir  of  llie  stomach. 

The  cDiixish  iifii  of  the  food  ajipears  to  have  little  iuHuenee  on  its  rate  oi 
diseharire  from  the  stonmch-   at   least  in  tin"  case  of  potatoes.     Dilution 

of  protein   f 1.  iiowever,  incicases  the  rate.     Distinctly  hard  i)article.s 

in   the   t'ood  retard  the  stonnn'h  e\acnalion. 

There  is  nsnally  a  considerai'le  annnnit  »(  i/iis  in  the  part  of  the  stomach 
aliove  the  entiance  of  the  caiilia,  on  accomit  of  which  this  part  of  tin 
stonnn'h  has  sometinn's  iieen  called  the  stcnnach  liladder.  In  the  uprij,ht 
position  this  ^as  forms  a  lirijrlit  area  in  the  x-ray  plate  ^  b'ij;.  I.m),  hni 
when  the  ])erson  rediin's  it  sjn-eads  to  a  new  location.  Its  prescnec  nia\ 
inllncncc  ;;astric  digestion  liy  ]>reven1in};  the  contact  of  the  food  with 
the  niucons  memhrane,  and  hy  interferini;  with  the  efficiency  of  the  peri- 
staltic waves  in  nmvinu  the  food.  ( •onsideralile  >;as  therefore  retards  the 
emptyin-r  of  the  stomai-h,  as  has  lieen  shown  experimentally  hy  x-ra.\ 
oh.servations  on  animals  fed  with  the  standard  anniunt  of  foml  followed 
h.\-  the  introdnction  of  air.  It  was  imted  that  the  air  did  iH)t  diminisli 
the  frcqnency  or  strength  of  the  i)cristaltie  waves,  but  that  those  eonhl 
not  efficiently  act  on  the  food.  When  alonir  with  sias  there  is  also  aton\ 
of  the  stonnich  walls,  the  retardati(ni  in  the  diseharire  will,  of  course,  he 
still  more  proiionnccd.  The  temperature  of  the  swallowed  food  does 
init  appear  to  have  mindi  inlhuMn-e  cm  the  stonnn-h  movements  ny  on  the 
the  rate  of  discharge  from  the  oi'izan. 


.    I 


'I 


,l\ 


fii.^'M: 


ii:li  J 


H 


TIIK  MKC'IF.*  "l"  Ms  'il'  I);  ;i%STI<»N   ((Vi)ird) 

THE  MOVEMEw  To  OF  THE  INTESTINES 

The  Icnirtli  «'f  llio  siiimH  iiitcsti.u'  aii.l  the  size  "f  Hio  cecum  of  the 
l,,r-,.  intesline  vary  (M.iisi.lcrahly  ii,  ditTeiTiit  animals.  In  the  carnivora, 
Mich  as  the  .-at,  the  small  intestine  is  relatively  short:  in  the  herhivora, 
nlativelv  lun-r.  Thus,  it  is  three  times  the  length  of  the  ho.ly  in  the  eat, 
i,i„l  four  to  six  times  in  the  (h)-,' :  whereas  in  the  ^'oat  an.l  sheep,  it  may 
l,e  nearly  thirty  times  the  h^nsrth  of  the  hody.  lii  the  earnivora  the 
.-..■um  iseither  absent  or  rudimentary,  whereas  in  those  herhivora  whieh 
,|o  not  have  a  divided  stomach  the  cecum  is  very  Inr-e  an,l  sacculated, 
;,s  is  also  the  colon.  The  reason  foi'  the  irreat  si/e  in  hei'hivora  is  that 
practically  the  whole  of  the  dijrestion  of  cellulose  takes  place  in  this 
part  of  the  srut.  This  dijiestion,  as  we  shall  sec  later,  does  not  depeiid 
nil  jiny  secn^tion  poured  forth  hy  the  animal  itself,  t.ut  up""  <!"'  "'''''»" 
nf  hacteria  and  of  certain  en/ymes  (cytases")  that  are  taken  ^vith  the 
xciTctahle   food. 

Movement  of  the  Small  Intestine 

The  movements  of  the  small  intestine  have  heeu  studied  (])  hy  the 
i.ismnlh  suhnitrate  and  x-ray  method,  i  "2 )  hy  ohservintr  them  after  o|.en- 
in-  the  abdomen  of  an  animal  snbmer^'ed  in  a  bath  of  physiolotric  saline 
;.t  bodv  temperature,  C?)  by  observing  the  .dumires  in  pressure  produced 
ill  a   tiiin-walled  rubber  balloon   inserted  in  the  lumen  of  the   -ut   and 

,.,,,n,ected  with   a   r )rdinc  tambour   ((•"isi.    HiOi.   and    i41    by   excisim; 

purlions  of  the  intesline  and  keepin-  them  alive  in  a  bath  of  saline  solu- 
tion at  body  tcmp.'rature,  throuiih  which  oxyiren  is  made  to  i>ass. 

Tin;  Si.(;Mr.\TiN<i  Movkmi.nts 

When  a  suitably  fed  animal  is  placed  on  the  holder  for  examination 
liy  the  x-ray  method,  no  movement  in  the  intestinal  .shadows  is  generally 
.  i.served  f.u'  sonu'  time.  The  first  movement  to  appear  is  the  breaking  of 
..lie  of  the  eolunnis  of  food  into  small  segments  of  nearly  eiiual  size. 
!•:;„},    of  these    segments   a-rain    (["'"''^'y    '''vides,    and    the    neigld)oring 


lives  suddeidv  unite  to  form   new  segme 


nts,  ami  so  on,  in  a  manner 


4(1: 


i 


■ 

1 

1 

I 

^ 

j 

1 

fl 

fi 

i 

,11 


■  H 


Kit 


hllll.STII>\ 


^! 


i  ■■  I 
i    .1 


I 


i;  ii 


i\ 


wllicll  will  III-  liiadf  fif.ll'  liy  cnllMlllilit;  V\ii.  llil.  Tllis  ilnilluillc  m  >; 
till  iiliilioii.  as  Ciiiitioii  has  rallt'il  it.  i'i>iitiiiii<-s  witlmiit  (M'ssalimi  I'm'  iiiuii 
tiian  hail'  an  iioiif,  and  tin'  t'dini  sliadnw  (h)«'s  iKit  niiMnwhilf  sci-ni  to  fliaii^" 
its  ]>(isiti(in  ill  th(>  aluloiin'ii  to  anv  cNtciit.  'I'lit'  splittiiiji'  up  of  the  sci; 
iiii'iil  and  the  nishiii;;'  lou'ctlicr  o1"  tlic  iicijflihoriii'!;  lialvcs  inocecd  as  a 
rule  with  irit'at   rapiditv:  tlius,  il'  uc  cuiiiit   tlii'  iiiiiiilur  nt'  dilTcrt'iit   si'li 


liw.     1'''        Ai'p.ir.ifi-     iMt      II  .  ..tiiiiii,'     tr,-ntt" 


nf     Ulf     iillr^linr  (  I'i'iTll      l.'lik^"li 


iiiriits  diii-inu  a  di'liiiitc  |n'iiiid,  wc  may  lliid  ihc  rate  of  division  in  tli 
cat  to  he  as  lii<.di  as  'Js  cir  .">ll  a  iiiinutt'.  In  man  the  divisions  occiii'  at 
t'ii'i|iii'iicy  ol"  aiipi(>\iiiiati'ly  IK  |i.t  ininntc.  wliich  coii-csponds  to  the  t'l' 
(|iirnry  with  which  suinids  can  lie  heard  when  ilic  alidoiiicn  is  auscultate. 
Altliounh  half  ail  lioiir  is  the  in'iiod  wliicli  this  process  usiiall;»  <■■ 
('ii[iies.  it  may  last  eoiwidiMaUly  Imiirer.  in  eert.iiii  animals,  .sucli  as  r: 
ralililt.      Ncniiieiiliiitr      lllineineiits      ha\e      llul      lieell      ohse|-Veil,      lillt      ilisle 


r  ( 

;i     I       j 


in:    Ml  (11  VNMSMS   (IF    DKilXTIdV 


4f.r 


,r  tliciii  a   I 


livtliiiii  ■  1(.-iiii(l-ri<i  sliiltiiiir  «il'  the   niiisscH  o 


I'  rcKiii  iiiiiiiL 


luiiicn  (• 


f  tlic  •lut.     .ii>i(lly  rcpcatf.l  U>v  niaiiy  imiuitcs 


Whoii  tlic  iiitcstiiK's  arc 


ll.iatcd  cut  in  a  warm  liatli 


liath  of  saliiif  siiliilion. 


it  is  seen  tliat   tlic  iliytlmiic  sfsriiifiitatioii   is 


caused  liv  narrow  riM)js  ( 


I    IS  seen    iM.u    MM     >>•.' ^ 1       1     t     «l 

.„ntraction.  In.lcr  Mich  conditions  also  it  is  oltcu  noted  tlat  tin 
l,„.,,s  of  inlcslinc  sway  from  side  to  side.  Tin.  l,all.,on  method  also  re- 
^e•,ls  tlic  presence  ,<f  sli-lit  waves  of  coulraclion  tliat  pass  rapidly  alon- 
,l,e  -nit  and  follow  ea.d.  otl.cr  at  the  rate  of  twelve  to  tliirtecn  per  minute^ 
l'.„th  ot'  tl.c  musclar  e.,ats  of  the  intestine  arc  involved,  and  u  is  believed 
,l,at  tl.c  ..(nitraetions  arc  rcsponsil.lc  n.,t  .ndy  for  the  pcndular  movc- 
„„,,ts  hut  lor  the  rhvthmie  seirmenlatio,,  ol.served  l.y  the  x-ray  method. 
Veeordinir  to   this   view    these   mov.mcnts   arc   eonstantly   passing   a  ..11- 

,1,..  intestine,  and  heco. xa-'cratcd  hy  the  mechanical  stimulus  which 

,.  oftVred  hv  the  masses  of  food  to  such  an  extent   that   they  divnle  the 
„„sscs  into  portions.      i'he  evidence   for  tiiis  hclief   rests  on   the   fact   that 


vli,.»   111,-  |.c.<iti"M  .i!  .livM.i'i  aii'I   "1    •   1-  -li">>"   "■'"    '""    "  •' 
.;iiiiiK   t"Kt-tlitr.      ilM-iiii    t";iiinuii.' 

When  the  contraction  is  studied  l.y  the  l.alloon  mctln.d,  it  l.cconics  ma  ked 
„vcr  the  mi.hlle  of  the  hallo..n,  uhcre  the  greatest  tension  exists. 

Se-eral  functions  can  l.c  assigned  to  these  movements.  They  cau.se 
intimate  mixture  of  the  f-.od  with  the  di..^"stivc  .juices,  and  l.y  hrinmn^- 
ivcr  new  p..rti..us  of  fond  in  contact  with  the  mucosa,  they  cnc(.ura.'c 
absorption.  Thev  also  have  an  important  massairin-  inlluencc  (.n  the 
Mood  ami  Ivmph  in  tlie  vessels  (.f  the  intestinal  walls.  Indeed,  the  pas- 
s;nrc  of  lymph  from  the  lacteals  int..  the  m.'scnteric  lymphatics  seems  t.. 
lepend  very  lars;cly  up.tn  th.'se  movements. 

Tin:    ri.KIST.M.TII'    M.tVKMlNTS 

The  oth.^r  m..veiiicnt  ..bM-rv..!  in  the  small  int.-stino  is  that  known  as  the 
l.nstalll,'  mwr.  It  occurs  in  tw..  f..rms:  (1)  as  a  sb.wly  a.lvaneing  e.m- 
iraction  (I  to  '1  em    per  minutc>.  piv.-e.lcd  by  an  inhi.  .ti..u  ..t  the  walls 


1  prneeedins  only  throii'^'h  a  sln.rt  .listauce  in  a 


aid  pr 


il  (4  to  r»  cm 


) :  an 


A*^^ 


TT 

'iW 

'     : 

1,! 

II 

i':    ij 

m 

iiM 

m\: 


M\(\ 


i>i()i:sTi()\ 


I '_' >  i\s  ,-1  swilt   iii(i\  ciiit'iil  cnllril  ilic  iiinnldllii    nisli.  wliich  s        jis  with 
out  piiiiM'  I'lir  iiiiicli  loiitrci'  (list;iiiiT>  jilmiy  llic  (■■•iiial. 

I-'in'tlicr  .iiijilysis  .it'  tin-  |i(  i  isl.iltic  w.nc  <mii  icinlily  hv  w,iilv  by  the 
liallcMPii  iiictlHid  I  I'ii;.  lti"J  .  1 1  tlif  (iul  is  piiiclicd  jiliuvc  tlic  Imllooii,  a 
niai'k'il  iclaxaticiii  iicciirs  (i\(i'  it,  and  lliis  rrlaxatioii  extends  t'lir  alKHit 
t«ci  tVet  down  tile  intestine.  It',  nn  ttie  dtliei-  liand,  tile  j^ut  is  pillelied 
a  little  Im'Iiiw  the  situatiiiM  nl'  the  hallncm,  a  limir-eontinued  eniiti'actidii 
oc-eiiis  (i\  ci-  the  lattcf.  The  e(ineliisiini  that  x\  e  may  di'aw  fi'oiii  this  result 
is  that  tlie  stiiiinlatiun  nl  tiie  u'llt  eailses  eonl  laetion  ahuve  the  |)()iiit  ol' 
tlie  stiiiHiliis  and  relaxation  lieluw,  tliis  heinu-  known  as  "the  law  of  the 
intestiin'"- -(  Hayliss  and  StaiTin<;i.  We  ha\c  seen  tiiat  it  applies  also  in 
tlie  ease  of  the  cardiac  and  pyloric  sphini'ters. 


.^1 

■ 

.1- 

I 

w 

\ 

r- 

1 

Mj' 

i 

m 

._'.  lilt, -till, ,1  ..iiilr:iil!.ii«  'l..-ill.,.iii  lin-lll.,.li  al'tn  i^.i-ii 
lintii  \.i^i.  Ml  .  Ii.'iiii>;4!  .iiiMil.itii.il  ^ihi.vi'  III  :,n.|  l.<l.i> 
■  ■iilr.ii  lii.Ti    :i'ii.ti\ilv         il'i'.ui    St.-iiliii^'  1 


'it    the   iitiiliitiiiiiiil   K.-mvliu   anil 
'I    the   ti.-illi.i.n    c.itis,s    ril.iva 


Tin:  I'liYsiui.dcn-  N'ATrur,  ok  tiii;  FJiivtiimii-  .\m>  Pi.itisT.vr.Tir  1Iovi:mi:nts 

Interestintr  inforination  in  this  connection  has  heen  trained  l)y  ohser- 
vjition  of  the  iiclia\  ior  of  the  nio\t  iiients  after  the  aiiplicatimi  of  dnisrs 
to  the  ;.'iit  or  after  cnttiny;  the  nerve  sui>ply.  The  rhythiiiie  movements 
are  not  atVectecl  liy  the  api)lication  of  nicotine  or  cocaine.  .Since  these 
drujis  ])araly/.e  iiervtJiis  stiiictiircs  it  has  heen  conchnled  tl'.at  the  rliythmic 
moveiiients  are  iri.\oi.'enic  in  oris:in.  The  (piestiun  is  not  a  settled  one. 
however,  for  it  has  heen  fonml  liy  .Atafiiiiis  that,  although  strips  of  the 
lonuittidiiiid  muscle,  isolated  in  oxytrenatcil  saline  solution,  will  continue 
to  heat,  they  (h)  not  do  so  if  the  adhcri'nt  Auerhach's  plexus  of  nerves 
is  strii)ped  olT  from  them,  'i'lie  nature  of  the  ))eristaltic  conti'actions  i^ 
more  definite;  they  must  clearly  depend  upon  a  local  nervous  sti'Uc- 
tore,  since  they  are  paraly/ed  liy  the  application  to  the  srut  of  cocaine  or 
nicotine.  This  local  nervous  system  no  (hiulit  also  resides  in  Auerbaeh"- 
plexus,  which  nuist  therefore  he  considered  as  conipley  "uoufrli  to  he   fscc 


Tlir    Mi;i  IIAMSMS   UK    l>l(ii;STIO\ 


4G7 


|,,i^r,.  7<»ti)  cnilnw.'d  with  the  |M>\\.-r  ni'  liiri'ctiii!.'  iktvcmis  iimmlscs  so  us  to 
liiin-r  alHiiil  rcliixaticMi  nf  ilic  v'Ul  in  front  of  ilic  stimnlus  and  eoiitrac- 
lion  ovof  it. 

Ni;i!V(>rs  CoNiudi.  uK  Movk.mknts 

Tlic  intliicncc  of  tlic  rcntinl  nervous  s\  stci tlic  iiitfstinal  inovi'int'tits 

li.is  litH'ii  stn<li«'<l  liy  the  nsiiai  nn'tlio<ls  of  cutliiiy:  and  sf iinulatini,'  tlio 
,  \trinsic  niTVf  siipiilv,  Tlirou-ili  tlic  s])lan<'iinic  nerves  tonie  iithibitorii 
impulses  are  conveyed  to  the  intestine  i  except  tlu'  ih'ocolic  sphincter), 
lor  after  tliesc  nerves  are  severed  tlic  luovciiieiits  hceoiiic  iiiorc  distinct, 
iiidccd.  in  iiiaiiy  animals  after  opciiinj;  the  aliiloincn  no  intestinal  movc- 
iiKiit  can  1m'  ohservcd  until  tliesc  nerves  have  liccii  cut.  Stimulation  of  the 
|i(  rijihcral  end  of  the  nerve  also  inhibits  any  movement  which  may  mcan- 
uhile  lie  in  i)ro<;rcss.    Tlic  iminilses  through  the  vajius  nerve  arc  of  an 


.1      The  tllict  of  f\cilati(in  of  b"th  siiLiikIiiik-   lurvis 

StarliiiK) 


III   llic   intc^tii.al  coiitraftioiw 


nppositc  cli-aractcr.     Section  of  these  nerves  has  little  etVcct.  luit  .stimula- 
lion  causes  contraction.     (Fi<.'s.  Hi:!  and  104. ) 

i>y  ohscrviiiK  the  rhythmic  contractions  of  an  isolated  strip  of  the  small 
iMtestinc  suspended  in  a  hath  of  oxy-reiiatcil  saline  solution  at  Imdy  tem- 
perature, it  can  readily  lie  shown  that  the  presence  of  even  a  minute  trace 
nf  epinephrine  is  sufticient  to  produce  c<miplcte  inhihitioii  of  the  movement. 
Tlie  i)arallclism  l.etween  the  elTccts  of  splanchnic  stimulation  and  those  of 
•  piiicphrinc  injection  is  very  sisriiificaiit,  for  in  this  way  the  marked  inhi- 
Mtion  of  intestinal  movement  whicli  occurs  durin-r  frisilit  may  possibly 
'<•>■  explained    (see   pa^re  T^i'il. 

Tho  circular  muscular  coat  of  the  last   two  or  three  centimcnters  of 

itie  ileum  liefove  it  joins  the  ccciim  is  definitely  thicker  than  the  rest  of 

'his  coat,  indicatiuir  that   it  has  a  sphinctcrdikc   action.     This   iUocoUc 

phincfrr,  as  it  is  called,  opens  when  food  is  pressed  a<raiiist  it  from  the 

ilonm.  hut  remains  closed  when  food  is  pressed  asrainst  it  from  the  cecum 


!       I 


!      * 


Maiii 


4riM 


ii|(ii:sTio\- 


It  I  lu'ifrmc  'ilpfvs  llic  l.iw  (if  llic  iiilcstiiic.  'rii;il  il  is  |)liysi(il()H;ii';i!l.\ 
ilistiiict  rioin  tlif  inus<'iil;itiiii'  ul'  llic  icst  of  liic  ilfiiiii  is  iiidicatfil  liy  lli' 
I'ac't  tliut  llic  siilaucliiiic  fiiiil  vairiis  iicivcs  do  not  alTcct  it  in  the  saiin 
way;  thus,  stiiniilation  ul'  tlic  s|>lMnclini('  causes  a  stiimjj  contiactioii  ni 
flic  s|iliiiiclcr,  wlicicas  it  is  iiiialTcclccl  liy  stimulation  uf  tlic  vai;iis. 

I'ci'istalsis  is  iiiudi  iiinic  I'ajiid  in  tlic  iIiiihIciiuiii  tliaii  in  ntlicr  ])arts  ni 
tlic  siii.-.ll  iiilcstiiic.  nuiiiiL;  the  litst  slaves  cif  diircsticin,  tlic  I'mid  (Miii- 
n.'irily  lies  mainly  in  tlic  I'ijjiit  lialt  nl'  the  alidipineii,  and  later  in  tlic  lett 
lialf.  Tlieic  is  coiisidciahle  \aiiati(iii  in  tlic  time  tlial  elapses  licfiiie  il 
enters  the  cdli'ii.  In  the  eat,  eariicihydiates  reach  this  jial't  ol'  the  jriil  in 
alioiit  fiiiii'  hours. 


it-f! 


0^:' 


i  i "  ■ '  I 


:1 


i; 


Movements  of  the  Large  Intestine 

On  acccilint  of  the  iri'fit  dilVeicnecs  whieji  we  have  already  scon  t" 
exist  ill  tlic  '..a'  and  relative  iiM|)oitance  of  the  colon  as  a  dificstive  orsan 
in  dilTcrciit  classes  of  aniniMls,  it  is  not  surpri.- iuj;  that  the  iiioxcnicnl^ 
observed  arc  very  ditVereiit  Mi'cordin<r  to  the  dietetic  haliits  of  the  aiiiiiuil 
A])paicntly  the  moveiiicnts  are  much  the  same  in  the  cat  as  in  man.  .\- 
the  food  jiasses  tlironsih  the  ileocolic  s)ihiiii'ter  into  the  eccniii  aii'l 
accumulates  then,  it  s'"i'liially  sets  up,  h\  its  |)ressurc,  a  coiilractioii  oi 
the  iiiuscul;ir  walls  of  the  -jut  soiiiewherc  almuf  the  .junetion  lietwcen 
tlie  aseciidiii<_''  and  transverse  colon.  This  wave  of  coHtiactioH  ihei; 
liesriiis  to  travel  slowly  toward  the  cecum,  without,  however,  l)ein<r  pre 
ceded  hy  any  relaxation  of  the  wall  of  the  siit,  ;is  is  the  cisc  with  ;i  tin. 


I'm;  Ml  riiASisMs  111'  iii(;i>i'i(>N 


W.) 


IMiislaltic  wavi'.  Tliis  liist  \vavc  is  somi  fullnwi'il  liy  otlicis.  with  tlir 
K'Miit  that  tlic  fnoil  is  t'drccd  up  into  tin-  <'cciiim,  against  tlic  liliiid  cud 
n(  wliicli  it  's  cntwdt'd.  liciiiir  iiifaiiwliilc  prcvi'iilcd  from  passii'ij  iii1<i 
liic  ilciiiii  liy  till'  opci-iitiou  of  tlic  ileocolic  siiliinctor  and  by  the  oldicpic 
iii.imicr  ill  wliicli  the  ilcimi  opens  into  the  ceciini. 

\s  the  fcsult  of  the  distention  of  the  ( mil  set   ii))  l>y  these  so-p.-Hed 

■11 V 


s  o(  casioii 


.inliiieristaltic  waxes,  a  true  coordiiia1e<l  iiciistallic  wa^c  i 
initiated,  aiul  |)asses  aloni;  the  ascending;  colon  preceded  liy  the  usual 
wave  of  inhiliition.  These  waves,  ho'vever,  disappear  hefcne  they  rpach 
the  end  of  the  colon,  so  tiiat  the   t'ood  is  aL'ain  driven  hack  liy  the  so- 


1-1 


ria     1'.?.     liLWi 


tunc      It      l.ikt  -      iM!      ,i 


Mill    I*'    ri  ai  li    llii-    \.iii"M<    I'Uil' 


of    llic    liirRe    iiitc^lii" 


■  ailed  antiperistaltic  x\  a .  cs.  The  etTcct  of  the  uioveiiients  is  to  knead 
and  mix  the  intestinal  contents,  and  thus  eiicoura<_'e  the  alisorption  of 
water  from  tliein.  The  resultiiijr  more  -olid  portions  then  eoUoet  toward 
the  splenic  tlexiire.  and  liccome  sei)arated  from  the  rcmainiiiH;  more  fluid 
|iiirtion  liy  transverse  waxes  of  constriction,  which  dcxelop  into  peri- 
staltic waves  carix  iiiii  the  harder  m;isses  into  the  distal  portions  of  the 
loloii,  xvhcie  they  colli'cl  cliicHy  in  tlic  sii;iiioid  tlexure.  The  desecndin;.' 
'•iiloii  itself  is  nexer  distended  xxilli  contents  and  mercl.x  serves  as  a  tube 
I'or  transferrinu  the  masses  from  the  Iraiisverse  colon  to  the  sii;nioiil 
l!c\iiie.  The  time  taken  for  a  capsule  of  liismuth  to  reach  the  various 
narts  of  the  lait.'-e  intestine  is  sl!"X'.  n  in  [  iir.   l'>-">. 

.\fter  a  eeitaiii   iiia>s  has  collected   in  the  si-rmoid  tlexure  and  rectum, 
ilie  increasing  distention  causes  a  rollex  exacnatioti  of  this  jiortiou  of  the 


470 


KKiKSTiON 


n,\, 


KUt  tliniuuli  cfiitiTs  located  in  the  spinal  cold.  The  impulses  from  tlies. 
centers,  besides  eontrnctintr  the  fcetiiin,  etc.,  also  coofdinate  the  coiitrac 
tioii  ot'  the  alidoininal  iiuiscles  and  the  relaxation  of  the  sjihineter  an 
so  as  to  brint'  about  the  ai-l  of  ilefecation.  Hy  the  skiaiiraphic  method  ii 
has  been  found  that  the  pcl\  ic  colon  irfadnall.v  becomes  tilled  with  fccc^ 
from  below  upwi'iii,  and  that  the  rectum  lemains  empty  until  just  befon 
defecation. 

Ekki:(  r  ok  f'r.iNirM.  rnNoiTioNs  (i\  Tin:  Movi.mknts 

Observations  of  practical  value  have  lieen  made  on  the  behavior  of  th' 
peristaltic  \vave  afler  various  intestinal  operations.  Afhr  an  iiul-lo-i  ml 
tniiiKtoniiisi.i  of  the  <.'ut.  no  evichnce  can  be  obtained  by  the  x  ray  methiHl 

that  any  hesitation  urs  in  the  movement  of  the  shadows  at  the  aiia^ 

tomosis.     On    the  other  hand,   when   a   lalinil  inidshiiiiiisl.t   is  established. 

stjijriiatiou   of  the   food   in    the   re!:ion   of  the  junction   iiiiiy   iir.   thiv 

liaviiiir  been    fouml,  on   o|tcinii!.'  the   trut.   to   I Jiused   by   the   accuiiiii 

lation  of  liair  jind  umliirestcil  detritus  at  the  oi»eiiin'r  between  the  op 
jiosed  loops.  Another  oltjection  to  lateral  anastomosis  is  the  fjict  thai 
in  perforiiiinu'  the  o|)ei'ation  a  consideiiible  amount  of  tlie  circular  inuscli 
is  cut.  whicli  interferes  with  i>iristaltic  activity.  ^loieovei-,  the  end  oi' 
the  jiroxinial  loo|)  beyoiicl  the  openiiiir  is  in  danu'cr  of  becomiiifr  filled  up 
with  h.-irdened  mateiial,  and  the  end  of  the  distal  loon  may  becoire 
invairiiiatcd   and    induce   obstruction    in    the   re<.'ion    of  the   anastomosis 

01)servations   have   .also   been    made   by    the   x-r;iy    melhod    on    the   be 
havior  of  the  intestinal  contents  fojlowinir  liiti xflnnl  nhxlnirtiini.     !t  has 
been  observe<l  that,  as  the  material   collects  in   the   uut   just   above  tin 
obstruction,  stronu'  i>erislalti<'   waves  are  set   u)),  which   move  the   food 
toward  the  obstruction  so  jiowerfully  as  to  cause  the  walls  of  the  canal 
in    front    to   become  bnlsi-ed,   until    ;it    last    the   ]H'essure   causes   the   coii 
tents  to  he  scpiirted  ba<d\  throii-jh  the  advancini;  rin;;  of  peristaltic  coii 
trnetioii.     These   waves   were   observed   to   succeed    one   another   rai)iilly 
When  n  portion  of  irut  is  icversed  in  jiosition,  the  jieristaltic  waves  eon 
tinue  to  travel  in  their  (>ld  direetiim  toward  the  duodenum.    The  efTect  of 
this   is  to  produce  a   partial    obstructiini   at    the   up]).T   end   of  the   re- 
eeptivo  gut. 

The  type  of  peristalsis  known  as  the  pirixtnllir  ruxh  i-aii  be  induced 
experimentally  in  animals  by  intravenous  injection  of  erirot.  It  prob- 
ably also  occurs  in  conditions  of  abnormal  iri'itation  of  the  srnt  in  man. 
nn<l  is  believed  to  be  the  (diaracteristic  activity  of  the  trut  after  a 
stroncr  purtre. 


.■■  I 


i'M 


li 


ClIAITKK  IJV 


lirXCKR  AM)  AITHTITK 

Kuntrcr  ntul  ai)i>ftiti'  ace  distiiid  si'iisatimis,  tlic  I'liriiicr  lii'iii!.'  (Idiiiiti'ly 
iiiiTclatod  with  cDiitractioii  of  the  fiii|)ty  sldiiiacli,  and  llii-  latter  bciiiK 
,1   ('(implex   of  seiisatiidis   diieratiiij;   in 

•iuli 


the    iieivdus   svstein    aiontr   w 


ith 


)t' 


Utah 


food. 


.\|)|)etite    is    thei'efol'e 


lie  tiKid 
1  hiiriil.N  e(iiii|ilex  nervdiis  iiiteu'iatimi,  \\hei'eas 
liuiiirer  is  a  nuieli  simplef  pniccss.  Ii  is  iiartieuhirly  with  hunger  that 
\yv  shall  ciineeni  (lurselves  at   picseiit. 

When  a  thiii-xvalied  iidihei-  hallooii  nf  piopei-  si/.e  is  placed  in  the 
stciiiiaeh  and  connected  liy  a  rnlilier  liiiie  with  a  walei',  liidnmt'iifni  or 
cliloi-ofofiii  iiiaMoMieter  I  made  of  w  idi'  irlass  tnliinfi  I..")  ein.  in  diameter 
and  iirovided  with  a  snitalile  lloat  im  tlie  free  iimli)  a  tracini;  may  he 
taken  of  the  movements  of  the  stomach.  I-'or  nse  on  man  the  capacity  of 
tlie  lialhion  should  he  from  T-'i  to  l.'il)  ciihic  centimeters.  The  record  thus 
uhtaiiied  when  the  halloon  is  placed  in  the  empty  stomach  of  a  innnial 
person  shows  four  types  of  «ave.  Two  of  these  may  lie  discounted, 
lieiii},'  duo  to  the  arterial  pnlse  and  the  respiratory  movements.  The 
third  is  known  as  the  taiius  rlniUmt,  and  is  eau.sed  hy  tonic  contractions 
of  the  fundus  of  the  stomach  of  varyinjr  amiilitudes  and  oeetirrins;  at  a 
late  which  varies  from  IS  to  'J'i  Jier  second.  The  lierioils  of  tonus  in- 
crease duriii};  tl  powerful  rhythmic  contraction  to  he  immediately 
descrihcd.  While  these  cliaii^'es  in  tone  arc  occurring,  no  suhjective  sen- 
'^iltion  of  hunsrer  is  exiieriein-cd.     (See  I-'i«:.   I(i7.~i 

The  fourtli  ami  nmst  siuniticant  type  consists  of  jxiwerful  rhijthmic 
riiiifracfioiis.  alternatinjr  with  periods  of  (|uiescence.  These  contrac- 
tions occupy  a  period  of  ahoiit  twenty  seconds,  and  arc  superimposed 
upon  the  tonus  rhythm.  They  >;radually  increase  in  amplitude  and  fre- 
'luency;  and.  in  the  case  of  younjr  and  vijiorous  persons,  may  jrradually 
pass  into  a  condition  of  incomi)lete  tetanus,  after  which  they  suddenly 
siihside,  leavinjr  only  a  faint  tonus  rhythm.  These  rhythmic  eontrae- 
lions  arc  definitely  associated  with  the  sensation  of  Inniirer,  and  are 
more  marked  the  more  intense  the  sensa'iioii   is.     Wlien   tetanus  oeetirs 


hu 


liLi'v   sens 


iitioii   is   eontinuiiiis.    hot    it    instantly    dis;ii>p(>ars 


Mhen 


the  tetanus  jrives  place  to  relaxation.     When  the  contractions  are  com- 
parativelv  feeble,  the  lensrth  of  the  jieriod  diirinp:  which  they  octur  is 


M     !! 


471 


472 


KKIISTKIN 


'i        i 


almiit  twelve  iiiiiiiiles.  Iieiiij;  Nlioiter  tliaii  when  |1 iilraetitiiis  iire  |Miw.r 

fill,   ill    wliieli   eiiM'   tlie    periiHls   are   always    initialed    liy    weakei iilraf 

tioiis  with  limit  iiit<'r\  eiiiiit;  |iaiises.  Filially,  the  iiaiises  liisappear  an. I 
the  eoiitraetions  lieemiie  iniire  ami  iimre  |>i'imi>nneeil  until,  as  above  men 
liiiiieil,  a  xirtiial  tetanus,  lastin!.'  I'roiii  two  to  live  minutes,  may  super 
\ene.  The  th;ratiim  of  the  hiinufi'  period  varies  from  one-half  to  one  aii'l 
a  half  iiours,  with  an  averaifo  of  from  thirty  1o  forty-live  iiiiiiiites,  ami 
the  numlier  of  individual  eontraetioi.s  in  a  i)eriod  varies  from  twenty  t.i 
seMiity.  I'.etween  the  huiiuer  periods,  intervals  of  from  one-lialf  I" 
tviii  iinci  one-hall'  Imins  of  ipiies'-enee  may  supervene.     (See   Via.   llih.  i 

.Similar  eont  rai'limis.  ofleii  passinj;  into  ineomplele  tetanus,  have  l>eeii 
oliserxcd  in  the  stomach  of  liealtliy  liil'niits,  some  (d'  the  oliservations  liav 
ill;:  heen   made  hefore   Hie   lirst    nnisiiii.'.     The   interviils  of   motor  ipiies 


riK.    1.,!,.       Di.iBr.Kii    ..t    Tli,ll,...|    l„r    I,,  ,,rlii.<   sl,„v.,.h    ii.mv,-  .1.  nl-        /(.    r-ll.l.i-t    I..1II.,...,    iti  sl..in.l.  |i 

',    kyinograi.li.      /.    ,  .iU    H..at    «itli    rir..r,iiiiK    rt.ig.      V.    m.iM..i.,.  I,  r,      /..    i„a,i.,i„rtrr   fliii.l  (I. mm" 

iHrin.    ,.hl.,r„f,„,„,    „r    «al.-n,       ■■!.    riil.l.rr    1..Ih-    .  nii,,,,  i,„k    I,.-,|I,.„„    «ril,    maii.imcler.      .S".  sl.niiK  li 
/      -i.lr    till.r    f..r    ii.flatinri    ..f    -tniTM,  h    hall".!.].       I  l-r..,,!    CarhoM   ) 


: 
1 

'  M 

'. 

i 

1 
■  1 

W'i '' 

1  \i 

i'i 

«/\ 

eenee  l»ctwt'oii  tlio  hiiiiirer  periods  jire  .shorter  than  in  adults.  In  ohser 
vations  made  during  sleej),  it  was  ohserved  that,  when  the  eontraetjons 
were  very  vigorous,  the  infant  would  show  si<;ns  of  restlessness  and 
miirht  awake  and  ery.  .\s  in  the  a<lult,  the  eontraetinns  are  evidently 
assoeiated  with  suh.ieetive  sensations  of  hnnu'e'-.  ( 'ontrnetions  cd'  the 
emi)ty  stomaeh  have  also  heen  recorded  on  a  lar-:e  variety  of  animals. 
iiieludiiiK  the  do<r.  ijdihit,  eat,  irninea  pi<r,  lord,  froi:  and  turtle.  The.\ 
vary  somewhat  in  type  in  dilTerent  animals. 

With  resrard  to  thi>  time  of  onset  «(  Hie  tonus  :ind  huntjer  ecuitraetions, 
it  has  hr-fii  observed  that  the  only  period  dnrinn  which  the  fundus  i> 
free  of  them  is  immediately  after  a  larsre  mejil.  After  a  moderate  meal 
the  tonus  rhythm  begins  to  appear  in  about  thirty  minutes.     It  graduallv 


HI   M,l  l(     \SI'    Xfl'l  TITI. 


»7:l 


iiiiMciisfs  ill  iiili'ii>ilv,  until  l)>   tin    tiiiK'  tlic  sldiiiiii'li  liiis  iH'iirly  ciiiplicil 

iM'lf    till'    tdllllH    lia>    Iprrnliii'    rMii^|iiclHMIs.    illlil     tllr    <.t  I'ulliri'f    hUlllfiT    ri)ll- 

iijiclioiis    iisiiiilly    lii'v'ili    to   «i>iM'iir.      SiipciiiniX'MMl    upon    llmsc    of   tlii' 
'..mis  rlivtlim,  Iniiitfi-r  paiiirs  may  «pp<iir  in 
(iiilrtius  tracfn  of  food. 


mil   till  1 1)  Ihr  stoiniiih  still 


*  "i 


-m 


]■■?.     Tta.iiig  of  the  to......  rli>tl..i' 


till    *f..ni;iih    f.i.;iii.    tl.ivc    liMi!"   aftt-r   a   m.  al. 


\\\  .stllilx  ill!.'  till'  sIi;h1(i\\  (.1'  till'  iilltlilii'  i)f  till'  >1niiiin-li  pii.ilui'i'cl  li.\ 
liM\iiis  n  porsoii  Of  iiiiiiiKil  swallow  two  lialloons,  one  insiijc  llip  othor 
.iiiil  with  a  juistc  of  liisniiitli  siilniitratc  ln'twiTii  tlicin,  it  lias  liccii  ob- 
Mi'vi'd   that    the    weaker   tyiH-   of   luiiisrer   emitraetioii    lie-jins   as   a    eoii- 


I  is,   168— Tracings  frnm  Iho  st.>Tnaih  .l.iritiK  Ih'-  riilt.iiiiali.m  iif  a  |.rri.iil  of  vignrnus  »M«tric   i.i.t.Kr. 
'.■..nlraciiriiii.      Oinhnlf    r-riginal    ?i?c.      (From    Carlson.) 


i  .'Mil 


^trii'tiiiii   iiivolviliir  the  i-anliae  iiid  of  llie  sloiiuii'li,  and   moving  toward 

t'lt'  i)ylni'ii'  end  as  a  raiiid  peristaltie  wave.     When  tlio  I'ontraelions  are 

iiy  viKOiiiiis,  this  wave  spirads  so  rapidly  o\oi'  the  strnnaeh  that   it  is 

lillieiilt  to  detpi-niiiiP  whether  it  really  i urs  as  a  very  rapid  peristalsis 

..rasa  eontraotioii  involvinjj;  the  fnndiis  as  a  wlioio.    Tiiesc  contraetions 


^iij 


474 


imtKSTION* 


i\ 


rcscniltlc  vtTv  cliiM'ly  tlu'  itiiivt'iiiriits  tliHl  liiivi-  »iiitiftiiiicH  liccii  oliHt'rvr'i 
nfttM- II  liisniulh  iiiriil,  and  wliicli  luivc  Ih'imi  tlioiiitiit  l>y  I'liiiical  nhscrviM- 
til  i.iilii'iiii'  II  li\  pfipri  isiiiKis  of  tin'  stitinai'li.  Tlu'  fiiiiiliis  in  tlu'ri'l'ipn 
not  I'litirrlx  passive  iliiiinu  ili(fi'<.tioii ;  for,  altlioiiuti  early  in  this  ar' 
tlirri'  Miii,\  III'  III)  i'\  iiji'iH'i'  of  I'oiitiiii'tioii,  \  ct  the  I'oht riirt ions  of  till-  tiiini- 
I'livtliiii  iiiav  appriir  aiiij  lircoiiii'  proiiouni'i'ij  lirfori'  tlit'  ><loMiai'li  is  I'li 
tiirl.v    riiipty.      Ill   other   wonls,   the  iliu'i'^lion  I'ontrai'tioiis   of  the   lillcrl 

sloiiiarh   ( SCI'  pairi'  4.'i|i    pa-^s  L'i'a<luall\    oNcr  into  tin-  hiiii«i't ntrarlion- 

of  tilt'  ciiiply  orjfiin. 

It  appiars  that  the  stoiiia>-li  i-ontractions  proilui-i-  the  hunger  si'ii>.,i 
tioiis  liy  caiislii);  stimulation  of  allVii'iit  nerve  enilin|;s  in  the  ninsrl< 
layers  of  the  viseiis.  Mefi'  pressure  on  the  iiiileosa  itself  iloes  not  ori^'inal' 
Kiieli  a  sensation:  thus,  suiliien  ilislention  of  the  luilloon  or  ruliliinu'  tli< 
iiineosa  uitli  the  elosed  end  of  a  lest  tiilie,  inserted  throinfh  a  UMsIri' 
fistula,  dill  not  eaiise  aiiv  sensation  of  liunirer,  unless  the  stiiiiulii.  \v,i^ 
so  stroiii;  as  to  eailse  a  eontraetioii  of  the  liiuseulatiire  of  the  sloniaeli 

It  has  lieeii  tlioiiirlit  liy  some  oliser\irs  that,  diiriii',;  hmiiier,  eoiitia'- 
tioiis  similar  to  Ihosr  of  the  stoiiiii  'i  also  oeeiir  in  the  lower  end  of  iIm 
esopha>rus.  It  i>  lieiiived  li\  ('aiUmi,  ho\\e\i'r,  lliat  these  eoiil  rael  ion - 
are  not  at  all  lespmisilile  for  the  hiiinrer  sensation,  iilthoiii»h  they  iii;r 
irive  rise  to  a  feelinir  that  soniethini;  has  stuek  in  the  esophajius.  Con 
traetioiis  of  the  intestine  have  also  heeu  ohserveil  in  huntrer,  Imt  it  is  (hmht 
fill  \\hetlier  they  have  an\thiiii.'  to  do  \\itli  the  eaiise  of  the  liiinni  i 
sensation. 

REMOTE  EFFECTS  OF  HUNGER  CONTRACTIONS 

It  is  well  known  that  durins.'  hunger  eertain  general  suhjeetive  s.\im|' 
loms  are   likely   to  lie  experieiieed,   such  as  a   feeliiii.'  of   weakness  and    i 
si'iise  of  emptiness,  with  a   tendency  to  headaehc    and   sometime,;  evei 
nausea    in    persons   who   are    prone    to   headaehp   as    a    result    of   toxemi. 
eonditions.     IFeadnehe  is  likely  to  lie  more  pronoiineed  or  jierhaps  oiil; 
])resent  in  the  mornins;  liefore  there  is  any  food  in  the  stomaeh.     Thi"-. 
symptoms  indicate  that  Ininjrer  eoiitraetioiis  are  associated  with  hyin' 
exeitaliility    of   the   central    nervous    system,    and    it    is   of   consideraM' 
interest  that   olijcctive  siirns  of  this  association  can   be  elicited.     If  tin 
kiioc-.jerk  lie  recorded  alonjj;  with  a  record  of  the  irastrie  contractions,  r 
will  lie  found  that    it    is  m,irkedl.\'  {'xa^'fii'i'iitcd  siiiiultaiicoiisly   with   IIm 
stroni;    liiiiijicr   contractions   of   the    emp1\-    stomaidi,    this    ailtrineiitalio 
liciti!^  fircatrst   at    the  !uii;lit    of  the  slotilaeh   eotii  Cif!  ions,    •.'.liiii    'lie   Ini: 
frer  pain,'s  are  most  intense,  and  fallins;  otT  airain  to  iioriiial  when  the-' 
disappear   (I'"if;.    I'iO).     I-'iirtiier  ehaii|j;es   occurring   during'  the   liun!,'n 


Ill   MilK    \\l>    M'I'I  Tl' 


47.'. 


ihtiihI  include  iiii  incrt'iiNc  in  the  piiUf  rnte  anil  Misodilnlntitui.  Uy 
'  niiiiKi*' n(»  |i|ftlivsniiiiriii|>tiir  Itai-iiiitN  nf  llii'  arm   miJumh'    i  set-  ;ia>fi'  '2'M)) 

iiiij  Ntiiinai-li  ciiiitraclioiis,  it  lia>.  Ih'cm  fiiuml  that  tiic  incicasi-  ni  \nlunic 
..iciiis  piiri  ixissii  with  till'  incri-iiMini;  tuniis  ol"  i1h>  Ntntuacii,  Init  tiiat  it 
liiiriiis  ti>  islirink  In  I'oi'i'  the  Htitiiiacli  fniitiai'tiiMi  has  icaclH'd  its  nia\iimim. 
I  •iM-asi(iiiall\ ,  hii\M'vi'i-,  as  in  ai-utc  liuiiircr,  a  sunnwhat  ilitTirrnt  rt-la- 
iiiinship  iiiilainM,  viisncimMtriftion  liriiii;  nnni'  piiiiuiiirnt.  hiirinu  each 
liiiHiftT    ciintiactiKn    IIhtc    in    aU<i    inncasnl    salivation,    tin-    (Iciri'cc    "I' 

vliii'h  vaiifs  with  ililTtitnt  indiv  iiliialn.  This  salivatimi  is  indfp«'nd('nt 
■  >\  ilic  laiHf  (Mipiitiis  "watcriiiK  ol'  thr  month  "  that  acr'umpanics  the 
ilioniflit    of  si^lit    of  apprti/.ini;    I'ood. 


r< 


>liJiiiM»frfiiniii»iiiiii>llll 

i      If.*>.       Sh'iw  Mij;    .nlMh,.  111. I 


1     111.     k.... 
»     (  '  .tti  I     It 


i)>iiiiiiiiii>i'iiiiiiii>»iniMiriiiiiiiiiiiilliiiii'»'i 

k     liiMi.  I      l'.i'iii>!i     'hiiiin;     111,      iiMlkf.l     hillH.ir 


:|,.,.  I      ,.,,>iii.M     .hi 

clM..ni    »', Ill-Ill 


HUNGER  DURING  STARVATION 

Dni-iii!,'   fiit'oiiMd    stiii\a1ion    for    loiii;    pcriocls    of    tinn'.    it    is    known 
'liat  lu'althv  individuals  at   tii-st  cxpiTicncf  intense  sensations  of  hnniier 

iiid  ajjpctite,  wliiidi  last  however  onlx    foi'  a   few  da\s,  then  I me  less 

iroiionneed  and  finally  aiiimst  disapi    i   •.     It   is  of  interest  to  know  the 

I  lationshii)    hetwcen    these    sensations    and    the    hmiu'er    contraetions    in 

'lie  stomach.     Tins  has  been  investiu;ated  liy  t'arlson  and  l.nekhardt,  who 

iiliintarily  snhjeeted   themseUfs  to  eomplete  starvatioi,,  exi fpt    for  the 

ikint;  of  water,  for  fonr  days.     Diiiinjr  a  irre.it  pa.'t  of  this  time  records 

t'  the  stoniacli  contractions   were  taken   liy  the  lialloon   method,  and  it 

WIS   found   that    the  tonus  of  the  stonnn-h   and   also  tin'   fre(|nency   and 

intensity  of  the  hiinirer  contractions  liecaine  pro-rro-dvcly  more  pronouneeij 

!s  starvation  (iroceecleij.     Towards  the  end  of  the  period  it  was  also  noted 

iiat   incimiplete  hiini;er  tetanns  made   its  appeaiain'c   w  lierc  ordinarily, 

IN  in   Carlson's  case,   this   t.vpe   of   hunger   eontiai'tion    was   infre(|in'iit. 

-iiisaf inns  nf  hnnsier  wen    prcsi'ti!    mnt'c  oi-  !e>s  thitrUijlioiit   tiu-  ]>rriod, 

'  injr  therefore  prohahly  due  to  the  pcrsistentl\    increased  tonus.     The 

liset  of  a  period  of  h\ni^er  contraction  could  usually  lie  foretold  hy  an 


»  "I- 


!!■;     I  i 


1 M 


'if'  ^if 


'.i  .■*; 


I    1 

I: 


47(i 


nKil.STION 


iiicicasc  ill  llic  liiiiiL-'ri'  si'iisiil  inn.  ;iiii|  .'is  llicsc  (Mint  rjict  ions  licciiiiu'  iiini  ■ 
iinirkt'il,  tlic  liiiiiirt'r  M'n>MtiiiiiN  iciimc  iiimc  intense.  On  tlie  last  i\i\\  «. 
staivatidii  a  Inii'iiinir  seiisat  imi  n  I'erreil  to  tin'  e])iirastiiiiiii  was  aclileil  !■ 
that  of  limijier.  Tlie  aippetite  ran  practieally  jiarallel  witli  tlic  seiis;i 
tioii  of  Iniiiirer,  and  liotli  of  these  sensations  lieeaiiH'  )terce])til)ly  iliiiiii 
ished  (III  tlic  fourth  or  hist  (la.v  of  stisrxation,  this  (liiiiiiiiition  heiiiL' 
how  (•\ ci',  most  mailed  in  the  sensation  of  i(|iiietit(>.  !n(lee(I,  instead  of  ai 
eau'eniess  for  food,  there  (|e\  ('jolted  on  the  last  day  a  distinct  repilsrnaiir. 
or  iiKJilTerence  towards  it.  .\ccoiii|)aiiyinLr  these  sensations  of  hiiiiu' ' 
and  a|i|)ctitc  a  distinct  iiiciital  depression  and  a  fcclini^  of  weakness  wci' 
cx|)('rieiiced  diiriii','  the  latter  jiart   of  the  stai'Mition   period. 

<  Ml  partakin;;  of  food  airaiii  the  liiiii'j'er  and  appetite  sensations  \(i\ 
rapidly  (Iisa|ii)eare(l,  and  also  i>iaclicall.\  all  of  the  tnental  depressioi 
and  a  irreat  part  of  the  feelinir  of  \M'akness.  Coiiipletc  rccovei'v  frmn 
the  latter,  however,  did  not  take  place  until  the  second  uv  third  d,i\ 
after  lii'eakinsr  the  last,  h'roiii  this  time  on  hoth  men  felt  nmisiial!; 
well:  indeed  they  stale  that  their  sense  of  wcll-|peiii<^  and  clearness  o: 
mind  and  their  sense  of  irood  healili  and  \iu''(M'  were  as  yrcath'  improxc'l 
as  they  would  have  hecn  \>y  a  iiionth's  vacation  in  the  mountains.  The; 
further  jxiiiit  out  tluit,  since  others  who  haxc  starved  I'or  huisrei'  ])erioiN 
of  time  uiuinimoiislv  attest  the  I'.ict  that,  after  the  lirsf  few  days,  tin 
sensations  of  hnnL'er  hecoiiie  Ics^  pidiiounced  and  finally  almost  dis 
.■ip|ie;ir,  they  must  li.ive  e\])ericiiced  the  most  distressiiii_'  period  diirin'j 
their  four  da\s  of  starvation.  .\lthoii'_di  the  huiiL'cr  sensation  w  ;r 
stroiitr  enoU!::h  to  cause  some  (lisc(niit'(nt,  it  could  liy  no  liM'niis  he  ealh': 
iiiarkc(l  jiain  or  sulTcrin'_',  .■iiid  \\as  at  no  time  of  siifficient  intensity  !■ 
interfere  serioiisl.v  with  work.  Mere  starvati(m  can  init  thoret'orc  !•■ 
desitrnated  as  acute  siilTcriirj'.  it  is  of  further  interest  to  note  that  dui 
iiiir  the  starxatioii  period  a  continuous  llow  of  secretion  of  acid  trastin 
Juice  was  found  to  occur  in  tiie  stom.ich,  the  in-csciice  of  this  acid  prol' 
ahly  cxplainiiiir  the  acid  (U'  liurniii'j  seiisati(ni  expeiieiieed  in  the  I'lti^.i^ 
triiim  on  the  last  days. 

CONTROL  OF  THE  HUNGER  MECHANISM 

Tlic  control  of  the  liuiiucr  mechanism,  like  that  of  any  other  mecli:ii 
ism  in  the  animal  hod v ,  mav  he  ell'ectcd  throiiirh  the  nervous  system  <■ 
it  mav  depend  on  the  prcseiice  of  chemical  sniistances  or  lioriiioiics  ; 
the  lilood.  .\s  a  matter  of  fad,  it  can  readily  he  shov\  n  that  both  tho 
methods  of  control  are  operative,  and  we  will  iiou  c(Hisi(|er  lirielly  son 
of   the    f,-icls   iijioii    ■.'.hicli    this  coiiciiisiiin   depends. 

Altlimi'.'h  manv    facts  are  now    knouii   with   reiiard   to  the  uervinis  co 


Ill   Ndl  U     \Mi     M'I'I  TITI 


477 


i,,,l  (if  the  1 liiM'  iiii'cliiiiiisni,   il    is  dil'liciilt    Iti  l>i(i-«'  tlii'sc  tusietlicr  in 

Midi  a  wav  as  lo  i'oiiiiiilato  a  siiii|il('  tlicory  wliicli  tits  in  with  all  tin- 
..liscrvcil  tads.  We  know  tiiat  tlic  stuniacli  possesses  in  itself  a  loeal 
iirivdus  iiieelianisin  liy  wlii.-li,  like  tiie  heart  i>r  intestine,  it  eaii  antti- 
iiialieally  peiforni  many  ot'  tiie  iiiiivetiieiits  wliieii  are  exhihited  in  tlie 
iiitac-t  animal.  'I'liese  Im-al  moveineiits  may,  luiwevei,  lie  cniisidoralily 
liillilenee.l  iiy  iiii|)nlses  ( iaiisiMit1c<l  to  the  stcniiaeli  aloiin;  the  va'j:iis  and 
splani'lmie  iieives.  We  have  liieierciie  to  seek  for  e\  idenee  iiidicatini; 
tliM  relative  imixiitanee  of  the  loeal  lu  I'vous  iiieehanism  in  the  slomaeh 
itself  and  of  the  iiiii)ulses  transmitted  to  this  oriran  by  the  extrinsic 
iirrves.  We  imist  then  seek  the  position  of  the  eeilter  which  |)ereeives 
the  sensation  of  hnnjrer. 

It  will  he  simjilest  to  eon>ider  first  the  elTeet  of  section  of  the  extrinsie 
■M-rves  in  oliser\  atioiis  made  on  lower  animals.  Sirllaii  <ij  II"  si>l<niflniif 
■iirr<s  increases  }raslric  tonus  and  augments  the  «:astrie  hiin^'er  eontrac- 
finns.  Section  of  hull;  runtis  mrns.  performed  of  course  helow  the  level 
i.f  the  lieart,  leaves  the  stomach  in  a  more  or  less  h.\  potonic  condition. 
The  tonus  is  not  entirely  aholislic'l:  it  \aiies  somewhat  from  day  to  day, 
mid  may  heeome  quite  iiroiiounced  e\cn  thoULrh  the  va^i  are  cut.  In 
this  hypotonic  state  the  huiiirer  eontract  ions  are  diminished  in  rate 
,;iid  rejiularity.  Section  of  Imtli  the  splaiichnii'  and  vairns  nerves  throws 
!lic  stonuKdi  into  a  pernnnient  hypotoiius.  except  in  )iidloni:ed  starva- 
tiuii,  when  hnniier  contractions  develop  th,-it  are  usually  of  vreat  ampli- 
tude and  with  particularly  loiii;  intervals  lietwe.'u  the  contractions. 
The  yeneral  conelnsioti  to  he  drawn  from  these  experiments  is  that. 
;ilt!iiiULdi  c(implet.ely  isolated  from  the  central  nervous  system,  lhe 
'-loiiiach  still  exhiliits  typical  hiin';cr  contractions,  which  must  therefor.- 
iic  essentially  dependent  upon  an  automatic  mechanism  in  the  stomach 
'amII  itself.  Over  this  mechanism,  extrinsic  nerve  impulses  have  meiely  a 
ii'L'ulatorv  control. 


;  f 


■'i  r    'H 


Variations  and  Inhibitions  of  the  Hunger  Contractions 

The  all'erent  stimuli  that   may  set   up  impulses  iiavelinu:  hv  the  extrin- 
sii-  nerves  to  the  stomach  are  coincvcd  Iiy  the  nerves  of  sense  or  are  of 

i.-vchic  oriu'in.     Siimulati f  the  u'listatory  end  orirans  in  the  month. 

IS  li.\  eliewinir  palatalile  food,  alwa.xs  causes  an  inhiliition  of  the  loiius 
:i!id  a  diminution  or  disappearance  of  the  hunirer  contractions.  Kveii  the 
liewiuir  of  inilitTcreiit  suhstances.  such  as  paraftln.  siifiices  to  produce 
listiuet  iiihihition.  unless  in  a  ease  in  which  the  I'ontraction  has  ])assed 
'ito  a  tetanus.  It  is  of  interest  that  sw  allow  ini;  moxenieiits,  in  the  ah- 
1  lice  of  any  food  suiistance  in  ilic  tiMUilJi.  are  sufliciciit  to  produce  ;i 
■Miisitdrv  inhiliition  .if  the  :.;ast  lie  tonus     a   receptive  relaxation   of  th.' 


:i  .:1i 


478 


nitlMSTlOV 


ir     I 


i-'    lu 


stdiiini'li,   ;is   it    liiis   liccM    iijitlv   cillcd.      Tlic   iliiiiiiiution    in    tonus   aii>! 

limiv't'T  (■iintriii'tioiis  in   iIicm-  viirioiis  wjivs  is  a tniiianiod   hy  a  (liiiiinii 

tiiih  in  the  linii<;('r'  ]>ains. 

AfTcrciit  ni'i'vc  stiniiilatimi  atTcctiiitr  tlio  Imiijrcr  cunt ra<'t ions  may  als.. 
orii:inati'  in  llir  slonnicli  uninmii  itself,  as  lias  l)t'on  shown  Ity  Carlson  m 
liis  ])atii'Mt  liy  introihicins:  llic  vaiioiis  substances  to  lie  tested  tlii'ouyl 
a  tulie  into  the  stouiacli.  A  jrlassful  of  cold  water  introduced  in  llii^ 
\\i\\  inliiliits  the  tonus  and  the  lunn:i'i'  cont i-ietions  fof  ffom  thfee  to  (i\i 
minutes  unless  these  are  severe,  this  iidtihition  liein>r  followed  by  U" 
au!;mentation  either  of  the  tonus  or  of  contractions.  Ice-cold  water  lia- 
a  jricater  elVect  tiian  watei'  at  liody  tem])<>rature.  This  result  is  sonic 
what  dilTereiit  from  that  A\liich  most  men  exi>crience  as  the  residt  oi 
drinUiii!,'  a  ylass  of  cold  water. 

Weak  aciils  of  streii«:ths  varyiii<r  up  to  that  found  jiresent  in  the 
iiastric  .juice  itself  ()..')  per  cent -cause  a  marked  iiihiliitioii  of  tin 
liiiniier  Mioveiiients.  Init  this  inliihitioii  does  not  persist  until  all  the  aciii 
has  escajied  from  the  stomaeh  or  liein  neutralized,  wliicli  explains  uli.\ 
hunu'er  contract'ins  should  still  uciiir  when  an  acid  seci-etiou  is  i>reseiii 
ill  the  stomach,  as  in  sliirx  ation.     \ormai   irastric  .juice   itself  jiroducc- 

an   iiiliiliition.   which   is  no  doulit  deiicndent    U| the  acid  which   it  con 

tains,  ami  it  is  pioh.ihle  thjit,  at  the  same  li that  it  leads  to  iidiiliition 

of  the  hiumci'  contractions,  the  acid  initiati-s  peristalsis  of  the  pyloric 
reirion   (see  jia-re  4."(:!  i .      We.ik  alkaline  solutions  ha\c  mo  frreatcr  I'fl'ect  on 
the  liun-rer  contractions  than   an   iM|ual    volume  of  water.     Weak  soln 
tioiis  of  local  anesthetics,  such  ;is  jiheiiol  or  cliloretone.  arc  without  efTecl 

^^  ith  re<.'ard  to  olrnhiiJir  In  n  nn/i s  interesting  results  were  ohtaincij 
Wine,  lieer,  liraiidy,  ;iiid  diluted  imre  alcohol  inhihit  hoth  tlie  tonus  ami 
the  contractions.  The  duration  of  this  iiiliiliition  varies  directly  witlt  tin 
(|uantity  of  the  lieveraj.'e  introduced  into  the  stomaeh  and  with  its  alco 
hoi  percent;i'_'e.  TIicnc  oliservations  jire  appai-ently  not  in  harmony  with 
the  experience  of  most  men  th;it  the  takin<r  of  alcoholic  l)evera<res  .serves 
to  awaken  or  incr<'ase  the  iippetitc.  the  difTerence  heinjr  no  douht  due  t" 
the  fact  that  aiiiietite  and  hnnt'cr  contractions  of  the  stomadi  are  noi 
dependent  on  each  other,  appetite  lieiiiir,  as  we  have  seen,  a  eomplex 
)isyeliic  affair,  whereas  the  Inmu'cr  contractions  depend  tijion  a  loca' 
iiieehanism   in   the  stomach   wall   itself. 

As  the  iiihihilion  jiroclin'cd  in  on(>  or  other  of  these  ways  passes  otT 
the  hunu'cr  contra<'tioiis  are  resumed  at  their  previous  intensity  and  no! 
in  an  auunienteil  form.  I'r-om  the  promptness  of  the  inliibition,  it  wouh; 
a|)pear  that  the  stomach  contractions  are  afTected,  not  reflexly  throufii 
the  central  nervous  system  or  liy  ciiantres  in  the  chemical  oompositioi 
of  the  lilood.  hut   hy  a  direct  action  on  the  neuromnseular  meehnnisin 


HlNfiKR    Wn   At'I'ITITi: 


479 


ill  the  stdiiincli  walls,  iiiid  it  is  important  tti  hear  in  mind  that  tlio 
inhihifory  cITccts  on  the  stomai-h  rontvactions  ot"  the  I'tniiUis  may  procciMl 
i|iiitf  indi'pcntlontly  of  the  ehannfs  in  the  ])ylorii',  rp|j;ion  that  are  con- 
ici-ned  with  the  mechanical  jjidcesscs  of  digestion.  After  one  or  both 
(if  the  extrinsic  nerves  of  the  stomach  were  severed  in  do<;s,  a  certain 
■  li'irree  of  inhibit  ion  couUl  still  lie  indnced  liy  the  above  methods,  indicat- 
iii'_'  that,  alttionjrh  section  of  tlx-  extrinsic  nerves  depresses  the  inhibitory 
11  ilex,  it  does  not  abolish  it. 

X'arions  miti<;ations  of  the  hunjrer  contractions  have  been  discovered. 
SiiioUiny:  has  thi.s  effect,  an<l  compression  of  the  abdomen  by  tisihtenin<; 
tlic  belt  also  inhibits  the  contractions  jirovided  they  are  not  of  marked 
intensity.  Coiisidei'able  muscnlar  exercise,  such  as  brisk  walkinj;:  or 
iiiiiiiinjr,  causes  inhibition,  which  usually  persists  until  after  the  exei'- 
i-ise  is  disi'ontinued.  When  tlie  tonus  and  cont inactions  return,  in  this 
r.isc,  they  seem  to  be  somewhat  more  pronounced.  Api)lication  of  cold 
t.i  the  surface  of  the  body—  as  by  ])lacin<r  an  ice  pack  on  the  abdomen 
I'l-  takini;  a  cold  douche,  i)roeedures  which  are  well-known  to  induce 
i'Micased  neuronuiscuiar  tonus,  in  ficncral-  causes  an  i..iiibitioii  of  the 
'.;,i>tric  tonus  and  liunticr  contractions,  the  dcirrce  of  which  is  rouszhly 
|iiii|portional  to  the  intdisiiy  of  the  stimulation.  There  is  certainly  never 
.III  increase  in  the  jrasttic  tonus  or  huiiircr  contractions.  Ff  such  stimula- 
'iiiii  is  maintained,  the  inhibitory  elTects  on  the  stomach  {gradually 
'liiiiinisli,  even   though   the   individual   be  shi\  ('i-iii'j  intensely. 

With  rt'siard  to  tin    mm   ('(iitirs  coiiciriKil  in  this(   phniomi  nil,  little 

!liat  is  definite  is  known.  The  seus<iry  nuclei  of  tlie  vat;us  lu-rve  in  the 

imdulla  must  be  considered  as  the  priii<ary  ]iun<;er  center,  and  through 

■Ills  c<  iter,  not   only  inlluences  affeetinsr  the  stomach  contractions,  but 

:!!so  those  associated  with  tlie  luiufier  sensations,  must  be  mediated.       It 

'  'iiild  appear  from  observations  on  the  hunner  behavior  of  decerebrate 

iiiiiiiials  tliat  there  can  be  no  himtrer  ecntei'  located  on  the  cerebral  cortex 

il^clf,  for  such   aninnils  exhibit   practically  the  same  liuii<;ei'  effects  as 

■  iiinal   animals.   It    is  interestinjt  to  note  that,  at   least   in   the  case  of 

'    i('relii-ate  iiijieous,  this  huniiei-  beliavior  entirely  disapi)ears  on  removal 

tlic  o])tie  thalami,  where  im|)ortant  nerve  centers  havin<r  to  do  with 

ill'  bodily  responses  of  the  animal  to  Inniircf  impulses  would  therefore 

ijiear  to  lie  located.  These  observations  sui)port  the  sufi'/estion  that 
^  IS  been  made  by  several  neurologists  that  the  sense  of  pain   is  located 

iiicwhere  in  the  tlialamie  retrion. 

t 'iiiicerning  the  inlluence  of  i)sychic  states,  Carlson  says  that  in  his 
II   ease   the    liunyrer   contractions   became    weaker   and    the    intervals 

I  ween  them  greater  when  he  was  sudth-nly  awakened  duriii};  liis 
'  st  and  saw  two  of  his  friends  partakinjj  at  his  bedside  of  a  "feast  of 


!     H 


I  hi 


480 


PIOKSTION 


piiitt'iliiiiiM'  stciik  witli  iiiiiniis,  polalucs,  jiikI  a  tciiiiato  salad."  T1m>m 
results  aro  no  doiilit  <lu('  to  local  inliiliitioii  (li'pt'ndcnt  iipoii  the  i)sy('Iiii' 
stvrctioii  of  appetite  f?astrie  jtiice.  When  no  such  juice  is  produee(|, 
the  sif^ht  and  smell  of  jjood  food  does  not  appear  to  .itTect  materiallx 
Ihe  huimer  -ontractions  of  the  stomach.  \o  iloulit  it  stimulates  tin 
appetite,  Imt  that,  as  we  have  seen,  is  a  i)s>ehie  alTaif. 


m 


("IIAl'TER  LV 

TIIK  HKKIIK.MKAL  PKOCKSSKS  OF  DKIKSTION 

In  a  book  (Icsifincd  pi'imarily  foi-  crmical  workers,  it  would  be  out  of 

pliice  to  enter  into  details  eoneerninsj  tbe  biochemieal  processes  taking 

lilacc  durinjr  the  di<resfivo  i)rocess.    There  is,  liowever,  a  certain  amount 

oi'  I'undaniental  knowledge  wlueli  it  is  essential  that  \vc  should  consider. 

In  tlie   first  ]ilace   it  sliould  bi'  Iiorne   in   mind   that    in   the  digestion   of 

i;irlioli.\-dratts    and    pT-oteiiis,    various    intermediate    stages    are    passed 

tlirough    !-"f()i-e  the  linal   alisoiption   i)i-oducts  are   formed.     The  higldy 

r, implex   molecule  of  which  ])rotein,   for  exami)le.   is  eom])osed,   is  first 

111'  all  broken  down  into  several  smaller  but  still  highly  comi)lex  mole- 

'  ules,  each  of  which  then  undergoes  furtiier  disruption,  until  idtimately 

Hic  amino  ai-i<ls  are  set    free.     Certain   eii/ynies,   such   as   1ry])sin,   can 

I'Miry  this  ]>rocess  from   the  bc-jinnin'r  tlirou<rh  the   ijreater  part  of  its 

lourse  witliout  the  assistance  of  other  cn/.Mues,  hut  in  tlic  natural  proc- 

'  ^s  (if  diL'cstion,  as  it  occurs  in  tlu!  irastrcintestinal  tract,  the  difTerent 

^liiU'es  of  the  disinplion  arc  cnnlrollcd  by  dilTerent  eii/.yun's.     One  enzyme 

iircpares  tlie  food   foi'  action  by  the  next.     This  intcrdeiienih'nee  of  the 

itions  of  the  cn/.\in<'s  demands  that    some   i)rovision   shoidd   be  made 

hereby  each  en/vine  is  secrctcil  at  the  projter  time;  that   is,  when  tlie 

odstiifT  lias  alreaily  been  jiicpared  for  its  action  b"  ihat  of  its  prede- 

■ '  Nsor.     Thus,  it  would  be  useless  .'iftcr  fo<id  is  taken  t-      the  uastrie  and 

i'liiereatic  juices  to  be  sei-reted  at   the  sjime  time.     Instead,  the  gastric 

■  lico  is  secreted  first,  and  the  i>anci'ealic  onl\    aftei'  the   fooil  has  been 

'icpared  for  its  action.     This  correlation  in   function  we  Iiave  already 

111  to  lie  depeiiib'Ut   lart-'ely  on  the  action  of  hoiiiioiics. 

DIGESTION  IN  THE  STOMACH 

The  srastrie  .juice  contains  two  iiu|>oi-tant  digestive  agencies:  (1)  the 
Mzyme,  pepsin,  aiul  (2)  hydrochloric  acid.  It  is  particularly  in  juices 
•creted  in  the  cardiac  end  of  the  stomach  that  these  two  substances  are 

);it!d  present;  towards  the  pyloric  end  tlie  liydroclibu-ic  ,icid  entirely 
lisappears,  aiuI  the  pepsin  content  becomes  distinctly  less. 

4S1 


i 


I 


if  I 

Wm  1 

i\Ms    IK 


i  <  i 


!|,H: 


Is 


■  1 


4K2 


i>i(ii:sTio.\' 


The  Functions  of  Hydrochloric  Acid 


Tlic  fiinclioiis  of  liydrocliloiic  acid  may  Iio  coiivoiiicntly  divided  iritn 
Iiiiysi(iliii;ii';il  iiiid  lii.HJi.Miiic.il.  Tiif  fdriiici-  I'linctiidis  have  to  do  witli 
tlif  control  of  tiic  iiiovcinciits  of  the  stomadi,  including  tlic  o|)ciiiiiL' 
r)f  tlic  pyloric  s|)liiiictcr,  and,  after  llip  ,liynic  has  entered  the  dnodenuiii. 
\\  itli  tlie  secretion  of  pancreatic  Jniee  and  l)ile.  The  liioeheinical  fnnctimi. 
are  concerned:  (1)  in  assistinf;  the  i)epsin  in  the  diirostion  of  i)ri)teiiis, 
Ci)  in  lirin^'inj.'  alioiit  a  certain  amount  of  inversion  of  <lisaceliarid<v. 
and  C?)  in  liavinir  an  antiseptic  action  on  the  stomach  contents.  |{c 
fiardini.'  the  last  mentioned  of  tliese  functions,  it  may  he  said  that  tln' 
chyme,  as  it  is  ejected  from  the  stomach,  is  usually  sterile,  althoufrh  it 
may  contain  si)ores  and  certain  liacteria  th;i(  are  protected  against  tin' 
dijiostive  au'encies  of  the  stomach.  This  i)roteetion  is  afforded  by  an 
outer  covering,'  of  a  chitinous  nature  (si)ores),  or,  .-is  in  the  case  of  tin 
luliercle  haeillus,  hy  a  coveriiif;  of  waxlike  material.  It  is  believed  tliiji 
persons  with  strictly  normal  di!,'estion  are  much  less  liable  to  infection 
liy  such  bacteria,  as  tiiose  of  typhoid  and  cholera,  than  persons  with  hvN 
active  <rastric  secretion.  When  tlie  acid  of  the  jrastric  juice  falls  lielow 
the  level  at  which  it  develops  an  antiseptic  action,  various  bacteria  anil 
yeasts  irrow  in  the  stomach  contents.  i)i-oduein}r  by  the  resultintr  fernien 
tation  irritatinu'  oriranie  acids  and  fiases.  It  is  under  these  condition- 
that  yeasts,  sarciiuc,  and  lactic  and  butyric  acid  bacilli  find  in  the  •rastri'^ 
contents  a  suitable  nidus  on  \vhi<'h  to  grow. 

Tin;  A.MoiNT  OF  Acin 

It  lias  Ion<r  been  known  that  considerable  variations  in  tlie  amount  of 
hydrochloric  acid  in  the  -.'astrie  juice  are  associated  with  symptoms  of 
indiirestion.  On  this  account  a  more  or  less  elaborate  technie  has  been 
ilcveloiied  for  the  imrpose  of  determining  the  amount  of  hydroehlori.- 
acid  in  the  };a.strie  contents.*  There  are  three  thinfis  in  connection  witi: 
this  activity  that  we  may  measure:  (1)  the  total  titrable  liydrochlori.' 
aci<l:  i2)  the  free  liy<lrocliloric  acid:  and  (.'51  the  actual  hydronen-ion 
con(«<'nt ration.  The  determination  of  the  total  available  ai-ids  is  mad. 
by  titratin'r  a  measured  (|uaiitity  of  gastric  juice  against  a  standaiM 
alkali,  using  ])henoli>hthalein  as  an  indicator.  By  this  method  about 
7;")  c.c,  of  decinornial  alkali  solution  are  re<|uired  to  neutralize  100  or 
of  ii.u'mal  gastric  juice.  The  determination  of  the  free  hydrochloric  aci-' 
is  made  by  using  sjiecial  indicators,  such  as  those  of  (iiinzberg  an.' 
Tiipfer,  which  change  cobu'  at  a  hydrogen-ion  concentration  of  aboni 
Kr  isee  page  21).     To  produce  this  hydrogen-ion  concentration,  a  con 


<lhn.|. 


\ir   f'Miii,!    in   any   v.iliiiiu- 


liuiiial  ili.itiii 


Till:    lillicllI'Mlrxl,    CltMCIssiS   oK    lilclsTKiV 


4»:] 


siilcraldt'  (|iiaiitit\  0,(1.")  pw  ccMt  di-  iiHUf  <>)"  ..n  (ii'y:iiiiic  acid  is  iiooos- 
sary,  whereas  it  rciiiiircs  itiily  a  trace  of  liydnicliloric  acid.  Normal 
Iniiiiaii  (.'astric  juii-e,  wiieii  titrated  witli  one  of  tliese  indicators,  nivos 
a  (ifjiire  wliicli  corresponds  1o  aiiont  f>,(i:{  \  liyilrocldoiic  ncitl  (see  i)atre 
Jl'l.  For  till'  acewiale  (li'tflllliliatioli  ot'  the  h> dronen-ioii  eoiieeiitratioii. 
il  is  iieci'ssMiy  to  use  the  >_'as-cliaili  liiethiid   i  si'c  |iaire  "J!)). 

When  gastric  Juice  is  collected  throuu'h  a  listula  fioiii  an  empty 
stiiniacli,  very  little  ditTereuce  will  lie  found  lietwcen  the  i'l'ce  hydro- 
cliloric  acid  and  tlie  total  acid;  that  is,  lii'twecn  the  results  olitained  1>y 
the  secrnid  and  the  first  ol'  the  methods  descrilied  aliove.  Tlds  is  liecatise 
in  such  juice  there  is  no  ori;anic  mattei'  ea))al)le  itf  coiidiiiiiii(;  with  the 
lixdrcxddoric  acid,  and  there  are  no  other  acids,  such  as  lactic  or  hutyric, 

which   miifht    lie   i>roducfd    liy    t'ermentat 've    pi esses.      The   ditTcrenee 

lictweeii   the  two   titrations,   lioucxcr,  heeonies  (|uite   marked   wlien   ino- 

icin  r 1  is  underu'oinir  diu'cstion  in  the  stomach,  liecause  at   its  ditTereiit 

vl.iires  of  di'^estion  |iroteiii  eomliines  with  iiicreasinir  (|nantities  of  the 
liydi'ochloric  aciil.  'I'he  patholoiric  condition  in  whi<'h  theie  is  most 
ilifiiiitely  a  diminution  of  the  hydrochloric  aci<l  is  cancer,  eithei-  of  the 
^tolnach  itself  oi'  occasionally  of  some  other  ))art  of  the  liod.v.  An  in- 
crease is  i)articnlaily  marked  in  uh'er  of  the  stomach.  A  considerahle 
\ai'  ,ion  in  hy<lrocliloric  ,ii'id  nia>  howexcr  lie  the  result  meiely  of  fune- 
tiiiiial   (neurotic)   conditions. 

'I'lii;  Sol  Hci:  OK  iiii:  Arn> 

A  (|uestion  that  has  pn/zled  pliysioloj.dsts  for  many  years  concerns  llir 
III! cliiuusiii  h>i  ifhicli  liiiilriiihliirli'  iukI  is  sirnliil.  The  ))ereenta!je  of 
iiydroiddoric  acid  in  tlie  u:astric  Juice  is  consideralily  aliove  that  at  ^vhicIl 
any  animal  c<'lls  can  live,  and  \  et  this  acid  is  secreted  liy  the  lininu' 
iiienihrane  of  tin-  stomacdi.  its  source  licinu,  of  course,  the  soiliuni 
chloride  of  tlie  Mood  jtljisma.  llow  then  do  the  cells  of  the  srastric 
'j-Jands  lnin<;  aliout  ihe  sciiai'ation  oi  this  powerful  acid  from  the  per- 
fectly neutral  hlood  i)lasma  ?  In  the  liist  place,  it  is  si":nilicant  that  the 
iiiucous  memlirane  of  the  stomach  contains  a  hiirhei'  iiercentaire  of 
'hloriiH'  than  the  averaiie  of  other  oiyaiis  and  tissues,  indieatiiifr  that  it 
lias  the  iiowcr  of  alislractiiiir  chlorine  from  the  blood.  The  excess  of 
'lilorine  in  the  mucosa  must,  unueover,  lie  hut  a  very  small  fra<'tion  of 
iliat  actually  sei'ieted  inio  the  the  irastric  Juice.  The  chlorine  content 
"f  the  mucosa  of  the  cardiac  end  is  consideralily  irreater  than  that  of  the 
liylorie.  These  facts  indicate  that  chlorine  is  jittiai'ted  liy  the  srastric 
ri!k_  )>Ht  Ili(-y  11iro\v  lin  Imlit  oti  the  ijuestioii  as  to  wiif-re  the  hydros 
ililorie  acid  is  really  formed.  Is  it  in  the  cells,  or  only  in  the  lumen  of 
tlic  inland  tulies.'    That  is  to  say,  is  it  formed  liefore  or  after  the  g.'istric 


I 


f«Ri 


!     i 


;i 


r  'rlli 


4S4 


mOKSTION 


i:"- 


'I     I- 


■^  ■l-i 


.juii'i'  lias  lifcii  sfci-ftcd  fiiiin  tlic  cells*  After  iiitraveruiiis  injecfioii  m 
solutions  of  potassiniii  I'erroeyaiiide  aiul  some  inert  salt  of  iron,  sueli  as 
one  of  the  scale  iiicparatinns,  examination  of  the  jjastrie  glands  has 
shown  that  the  jtrnssian  Mne  reaction,  which  recinires  tlie  presence  oi 
free  miiuMal  acid,  is  most  pronounced  in  certain  of  the  parietal  cells.  .\ 
considerjdile  amount  of  the  jnccipitate  is,  liowever,  also  visible  in  tin 
lumen  of  the  glands  and  in  the  str)niach  itself.  (Vrtain  observers  affirm 
that,  although  some  of  tlie  parictid  cells  nniy  take  the  stain,  llie  vast 
majority  of  them  do  not  do  so;  and,  moreover,  that  cells  incapable  oi 
forming  hydrochloric  acid  (e.g.,  of  the  liver)  may  also  become  staineil. 
and  that  the  pn'<'i]>itation  may  occur  in  the  blood  and  lymph. 

The  coiifusi(,  .  in  the  results  by  these  methods  iirompted  A.  B.  Xlaca! 
Inm"  and  Miss  M,  1*.  I-'it/u'crald  to  investigate  the  distribution  of  the 
chlorine  in  the  cells  by  a  microchemic.il  method,  in  which  the  cldoridi-. 
were  preci])it!ited  with  silver  nitrate  and  the  silver  chloride  then  reduced 
by  exposing  the  section  to  liirht.  It  was  foinid  that  both  kinds  of  gas 
trie-gland  cell,  chief  and  parietal,  but  particularly  tlu'  parietal,  gave  tlic 
chloride  reaction.  Using  as  a  stain  a  substance  ( CyaiuiMiie"!  which  reacts 
blue  with  acid  and  rod  with  alkali,  Harvev  and  Hensley,''  however,  aver 
that  the  secictioii  of  the  dands  is  i)racfic;illy  neutral  until  the  foveola  is 
reached,  whore  the  stain  becomes  blue,  indicating  an  acid  reaction 
This  seems  to  show  that  the  acid  is  not  really  secreted  by  the  cells  oi 
the  gastric  glaiul,  but  is  foiined  after  secretion. 

Aeco..in)g  to  the  latter  investigators,  the  ctilorine  is  secreted  by  tin' 
cells  into  the  fovea  as  some  weak  ehloiide,  such  as  ammonium  chloride 
or  it  may  be  as  an  ester.  Shortly  after  its  secretion  this  weak  chlovi-b 
undergoes  a  hydrolytic  or  other  dissociation,  duriuir  whicli  free  hy<lro 
chloric  acid  is  liberated  and  .-immoniii  oi'  some  other  weak  ))ase  set  free 
Of  these,  two  products  of  the  reaction  the  we;d;  base  is  real)sorbed  by 
the  gland  cells,  but  the  hydrochloric  ,icid  is  left  behind  because  the 
cells  are  impervious  to  it.  Tmlirect  evidence  in  support  of  this  view  i^ 
alTorded  by  eertiiin  other  instances  in  whicli  hydrocliloric  acid  is  i)ro. 
dueed  by  tlie  action  of  cells;  thus,  the  mold  reiiicillium  glaueum  when  it 
is  grown  in  a  medium  c<intaining  ammonium  chloride  absorbs  tlie  am- 
monia but  leaves  the  hydrochloric  jii'id.  The  hii:h  ))enetrating  power 
of  the  amnnniia  ion  in  pra.'tically  all  cells,  and  the  fact  that  the  mticosa 
of  the  stomach  contains  a  higlicr  i)erceiitaLre  of  ;mimoiiia  than  any  other 
tissue  in  the  body,  must  also  be  considered  as  circunistaiitial  evidence 
in  favor  of  this  view. 

Whatever  !)0  the  mechanism  by  v.liich  !iydvn<-1i)ni-ic  acid  is  prodnce.!. 
there  is  no  doubt  that  the"  epithelium  is  impenetrable  to  it.  When  the 
vitality  of  the  epithelium  becomes  lowered,  as  in  anemia  or  after  parti,.' 


THK  iiiixiiKMicAi,  rmin.ssiis  of  DKiKsriov 


485 


Hc'clusiou  of  tiic  arteries,  tlic  afid  may  penetrate  the  cells  and  cause 
ilisfestion  of  the  .stomach  walls.  Hyperacidity  may  on  this  account 
lii'coiiie  dangerous,  as  it  lowci's  the  resistance  of  the  cell. 

Till  (lificstive  action  of  Injdrochloru:  acid  is  closely  linked  with  that  of 
pi  psjn.  with  which  it  will,  fhcicfofc,  he  consiilcred. 


The  Action  of  Pepsin 


It 


t  IS  I'oiiinionly  l)elii'vcd  tlint  l)efnro  its  secretion  pepsin  exists  in  the 
iclls  of  tiie  (jasti'ic  trlands  as  zynioireri  t'lanules.  The  chief  evidence  for 
tliis  helief  appears  to  ]tc  that  after  considei'alile  activity  the  amount  of 
/viiiofren  frramiKs  in  thi-  ".'land  cells  is  found  to  lie  dc<-ich'dly  dimin- 
ished. By  such  an  hypothesis  it  is  easy  to  cxphiin  certain  interestinp 
results  eoneerninj;  the  effect  of  wealc  alkali  on  the  activities  of  extracts 
of  tlie  mucotis  niend)rane  of  tile  stomach.  Wlu-n  the  mucous  memhranc 
is  extracted  with  weak  acids,  tlie  extract  is  very  active  protcolytically. 
If  tliis  so-called  pc|)sin  solution  he  made  faintly  alkaline,  or  even  only 
ni'uti'ali/ed,  nnd  asrain  made  ;icid,  it  will  he  found  to  have  lost  much, 
if  not  all,  of  its  activity.  On  the  othei'  hand,  an  aqueous  extract  may  be 
it'iiclered  sli^ditly  alkaline  for  a  shoi't  time  and  still  display  its  digestive 
activity  on  subsequent  acidification.  The  extract  made  with  water  is 
therefore  much  more  I'csistant  toward  all<a]i  than  that  made  with  weak 
acid,  and  the  difTercn<'c  is  explained  on  the  s\ij)position  that  the  water.v 
extract  contains  pepsinoi;en,  whereas  th(>  acid  extract  contains  pepsin. 

Tt  is  believed  that  there  are  several  varieties  of  pepsin,  because  the 
I'Plimum  concentration  of  acicl  in  which  ])epsin  derived  from  the  stomachs 
nf  different  animals  acts  is  not  always  the  same.  I'epsin  of  the  dosr.  for 
iN.'imiile,  acts  best  in  a  h.vdrogen-ion  concentration  eorrespondiufj  to 
'liat  of  a  O.n.')  X.  hydrochloric  acid  solution,  whereas  that  of  the  human 
•-toinach  works  best  at  a  concentration  of  0.0:?  \.  Different  pepsin 
solutions  also  show  a  diffd'ence  with  reirard  to  the  oi>timum  tempera- 
ture at  which  they  act.  and  with  reuard  to  the  nature  of  the  protein 
■■'.  tiich  they  most  reailily  attack.  Thus,  the  pejisin  of  a  calf's  stomach 
ilisiests  casein  \ery  rapidly,  but  coattnlateil  eirjr  white  only  slowly, 
"luM'cas  the  pe|)sin  of  the  i)i<:'s  stomach  acts  on  both  these  proteins  at 
iiliout  the  same  rate. 

It  is  well  krn)wn  that  the  activity  of  i)epsin  can  proceed  oidy  in  the 
presence  of  acids,  but  this  action  of  a<'i<ls  does  not  appear  to  depend  on 
ttie  hydrogen-ion  concentration  alone,  for  when  e(|ual  (Hiantities  of  the 
■~.iiiie  pejisin  are  mixed  with  i|uantities  of  different  acids  so  that  the 
li^droL'en-ion  concentration  of  the  mixtures  is  uniform,  it  is  found  that 


iliiiestion  proceeds  most  laiiiilly  with  hydrochloric  acid  and  least  rapidl.v 
"  ith  sulphuric  acid.     The  SO,  ion  seems,  therefore,  to  be  unfavorable 


lb      I 
I     \ 


.1 1- 


48(; 


i)iGi:sTi()\' 


for  peptic  iiclixitio.     The  acid  smiis  1 Iiinc  nilli  the  pintciii  lu-l'im 

the  i>cpsiii  attacks  tlic  latter;  U>v,  it'  we  (Irst  cninliiiie  the  inntelii  \\itli 
ai'iil   anil    thin    wash   i\\\iiy   all    traces   ol'    free   aci.l,   the    pinteiii   can   In 

(Infested    in   a    neutral    pepsin   solntiim    without    the   lilicrntir.n   of  anv    fi 

acid. 

Thi-re   is   evidence   t.i   show    tlwit    pepsin    itself  also   lieecniies  ciiiilhiiied 

with  the  prcptein  during.'  the  diu'estive  process.     If  a  pii )f  protein  Micli 

as  lilirin  he  ininiersiMJ  in  a  solution  ,.f  pipsin  and  then  taken  out  and 
"asiied  thoroui:]d.\  to  u'et  rid  of  all  adherent  pepsin,  it  will  he  found,  ni, 
plaeinir  it    in   a   hydrochloric  acid  solution   of  tlie   proper  strength,  Ihal 

peptic   dii:estion   pi eeds.        Advantaixe   iini.v   he   taken    of   tins    fact    t.. 

sei)arate  )iepsin  from  a  solution,  hut  tlie  Itest  protein  to  use  for  this  pur 
|)ose  is  not  lil.riu  l.nt  elastin.  Hy  such  a  method  it  luis.  for  example, 
hccn  slniwn  that  there  is  sono'  pepsin  in  the  intestinal  contents,  proviiii; 
thus  that  when  the  chyme  passes  into  the  intestine,  tlw  ])epsin  is  not.  a- 
used  to  he  thoujrlit,  innuediately   killed  hy   tlie  proteolytic  en/yn>p. 


rSit 


H    ' 


T'HonicTs  OK  Pii-nc  l>i(ii:sTiov 

With  retrar.l  to  the  ))roducts  of  trastri.-  di'_'estion,  little  can  lie  said 
here.  The  lirst  product  is  a  metaprotein  known  as  acid  albiiinin  oi 
syntonin.  It  is  pn'cipitated  from  the  di-estion  mixture  hy  neutraliza 
tion.  The  ucNt  prr)duct  is  known  as  jirimary  |)roteose,  lieinir  preeipi 
tated  liy  li.ilf  satuiation  with  auMMonium  sulphate.  The  third  i)roduci 
IS  secondary  iiroteose,  produced  liy  complete  satuiation  w  itii  the  above 
reau'ciit:  and  after  all  these  bodies  have  been  separated  out,  thpro  it 
mains  in  solution  the  fourth  product  |ieptoiie  which  amonjr  othei' 
thinu's  is  eharactcri/e.l  by  the  fact  that  with  the  biuret  test  it  -.'ivos  not 
a  \iolet  but  a  rosepink  color. 

It    lias  often    been    I'laiiiied    thai    alony    with    these    |noducts   a    certain 
amount  of  free  amino  .-leids  may  also  appeiir  in  a   peptic  diu'f'stivo  mix 
tiire.      This,    however,    may    be    due    to    the    action    of  erepsin,    mIiIcIi    i- 
usually  |)reseiit  in  pepsin  preparations.     It   is  important  to  note  that  tie 
term  jiriiiinsi  is  a  un-neral  oiie,  and  that  thi'ie  are  probably  iiiaiiv  vnrirticv 

of  this  substam-e.  ditTiriiiL'   from    one  .•mother  ; rdiiiir   to   the    proteii 

from  which  the.v  are  derivei]. 

The  chaniic  produced  by  jiepsin  .-nul  hydrochloric  ai'id  is  of  the  n.-iturr 
of  an  hxclndysis,  tor  it  has  been  found  that  the  amount  of  hydrojren  and 
oxyu'cii  in  the  diirestive  inoducts  is  irn-ater  tiian  flint  in  the  original 
protein.  It  is  by  a  simib-ir  process  of  hvrln.lysis  that  the  oth.er  jirnfeolyti. 
enzymes,  sindi  as  p;increatiii  and  erepsin,  operate,  but  this  does  no! 
imply  that   the  exact  u'roupintr  that   is  split  jiparl   by  the  hydrolytie  pro<' 


Tin:  mix  iicMHAi,  i-k(M'(>8kh  or  khikstidn 


487 


iss  is  tlir  siiiiu"  for'  ciH'li  "f  tlifso  cii/viiu's.  Imlri-il,  flicro  ix  foiisidornbh' 
cviiU'iico  tluit  |>rj)siri  (Iocs  hilt,  liki-  flic  iitlicr  ciizviiH's,  lirriik  up  the  limtt 
rliiiiii  of  iiiiiino  iii-iilN  that  ni'i'  liiikcil  tnKt'tlii'i-  to  i-ompiist'  the  polypcp- 
tiilcN,  but  tlint  it  only  splits  the  bin  niob-cnlc  ol"  allminiii  or  ubtlxilin 
into  si'Vpfiil  lai'dt'  (fi'oiips,  rarli  of  wliirh  is  coiiiposcil  of  loiiif  nniino-nciil 
ciiniiis.  Its  nction  nppi'ars  to  br  aiialoKo\is  with  that  nf  ainxbiso  on 
-i.irch,  by   wliicJi,  it    will   bf   rciiiciiibcrril,  tlio  biw:   polysaci-hariib'   iiiolo- 

I'lilo  is  split    into  smaller  j)olys; haiiiji'  molcciilcs,   wliich   then   liccomo 

iiltai'kfii  by  the  dcxtriiiasf  and  split  into  disaci-hafiilo  iiiolcciilcs  fsor 
|iiilfr  (>.'>(().  Thr  cvidrhcr  in  sn|ipor1  of  this  \  irw  is:  (1)  that  pi-psin  is 
inialilc  to  difri'st  polyprptidrs.  and  C.M  tliat  it  is  alili-  to  di^ri'st  i-crtain 
IHcifcins  upon  wliii'h  iTipsin   i  sci-  pajri-  4!KM   lias  no  ai'tiim. 

Till'  hydrolytir  splittini;  of  lai'do  into  sniallei'  |)rotoin  iiiolcculi's,  like 
lliat  by  whiidi  thr  idiains  of  amino  acids  in  thr  polyi)optidos  arc  subsc- 
i|iiiMitly  broken   u|>,  consists  in   a   breakinir  of  amiiio-carboxyl  linkiiiirs 

MK'O)    I  sec  pa;,'!'  ri)tS).  with  thi iiseipicnl  liberation  of  a  laru'e  iiiini- 

lier  of  luiattached  nmino  (;rou|)s.  The  nniuber  of  these  I'l'ce  amino  irronps 
i-aii  be  determined  (|iiantitati\ely  by  the  formaldehyib'  titration  inethnd 
lit"  Siireiisen.*  Hy  this  method  it  can  be  slniwii  that  from  the  very  start 
of  iicptic  diirestioii  tlie  number  of  free  amino  (.'roups  increases,  and  pari 

liiissii  the  power  of  the  diu'estive  ]>roducts  t indiine  witli   free  hydro- 

I'liloric  acid.  Tiidccd,  when  the  experiments  are  (bnie  (piaiititjitively  and 
the  ibuestion  allowed  to  i)rocerd  for  a  considerable  time,  the  increase  in 
the  fiirmol  titration  is  practical'.\-  eipial  to  the  decrease  in  the  free  acids 
as  determined  by  the  (iiinsberi.'  reaircnt. 

The  rate  of  jieptic  ili<;estion  is  usually  estimated  by  the  law  of  Schiitz 
iml  norissow .  accordin<;  to  wliich  the  amount  of  coa<;u1ated  albumin 
tlNit  is  dijjesfed  in  a  ^lett's  tube  is  proporf ioinil  to  the  siiujire  rout  of  the 
riiiininit  of  pepsin. t 

The  ])cpsin  \vliich  leaves  the  stoniacli  in  the  chyme  is  not  all  destroyed 
ill  the  intestine,  as  was  at  one  time  lielievcd  to  be  the  case,  for,  as  we 
liavc  seen  above,  some  pepsin  can  be  detected  in  the  trastrointcstiinil  eoii- 
tciits.  A  ])art  of  the  ])epsin  may  be  absoiluu  into  the  blood  and  carried 
liack  to  the  fiMstiic  irlands  to  be  ii>ed  as-'aiii.  This  would  account  for  the 
|ircseiice  of  antipe])sin  in  the  blood,  and  also  for  the  |>reseiiee  of  ])e]isin 
ill  the  urine.  It  is  probable,  however,  that  most  of  the  i)epsiti  is  dc- 
Ntroyed  after  it  enters  the  intestine. 


*In   Ihis  method    the   basic   cliar.iitfr   of    ttie   amino   ai'i<U    is   <Ifstr<)>c(l    Ity    the    forni.-iMcliyttc.    sn 

'I'-'it   a   ItiKluT  dciircf  of   acidity    dcvrloiis   in    thr   it:i\tnrc.      Ily   tittfrniir'.inK   the    iiutfiscd   acolitv    hv 

"Ointi    wiili   alkaii,    an    (slmi.Ttc    is   <ililaiiu-d    nt    tin-    millitn-r    ot    ammo    (troiips.       (  Src    caKc    .S'y  *.  i 

tTlu-  amount  nf  co:ii;tiI.iIi-d   rgn  all<ninin  di^:tsti-d   is  ascrLiintd   li\    iiuasniiMK  tin-   linnlli   dttii-stiii 

■luay  from  thr  end  of  a  rednmn   nf  martdatrd  egg  wImii*  c'mtaii'CiI   in   a   «lass  tnltr   I  Melt's   method) 

■s'-  f..l.li,   V.  \\        .\m.  .lour.   Physiol..   rMl^  \iii,  44K. ) 


I 


i'fl 


«  i'S 


488 


l>ll)|i>TIO\ 


Is]    !  ;  i 


Clottin?  of  Milk  in  the  Stomach 

Mosi.lrs  i(H  pourr  ,.f  .!i,f,...liiitf  [.i„(,.i,i.  tlio  Kiistric  jiiiw  i„  nUo  cn.low,,! 
«ith  th.-  pn.piMty  of  rlutiinir  milk.  Tliis  nrfinri  in  .•..mmnnly  aftiil.ui,  ,i 

to  the  prrsrri f  (iiintli.r  rn/yiiit.  lM-si.|.>s  jiipsin,   n!\w,\y,rn>ni,i :  l,u 

in  m-.'nt  ycjirs  (•..iisi.|,.n,lil..  .•..ntniv.Tsy  lins  vnanl  nmmi.l  fho  ,pi,.sii„i 
ns  fo  whrtlMM-  p,.p>ii,  aii.I  roimin  an-  imt  tli.>  sniii..  tiling,     On,'  str..i.  - 

nreiimonf  in  favor  of  this  vi.u   j>  ihaf  nil  .liu'.-stiv.-  Jiii that  an-  cap.il.!. 

of  .liifpstitm'  pn.Inn  ran  nNo  clot  milk      In  any  cnso,  «hon  enstrio  pii.-, 
nrfs  on  milk,  it  splits  fl.f  ••ascin*  of  tin-  milk  info  two  portions,  on.-  ... 
nhi<'h,  cnllfil  panwasrin,  immorliafrlv  conil.inrs  \(itli  rnl.'inm  to  form  iv, 
insolnMo  .'olloi.jal  componn.I.  mIuHi  is  pn-.-ipitat.,!  an.l.  In-  ••ntnnixliiM.. 
tlio  faf  of  tl...  milk,  forms  tlic  ••lot;  ttw  otli.T  proL-in  rrmiiins  in  soliHi,,. 
mul  \H  known  as  wlicy  all.nmos.-.     |-'r„m  sfn.li.-s  on  molecular  w.-iulit  i' 
is  iH'lirvp.l  tliat  tlu'  j.nra.'as.Mn  is  pro.ln.-r.l  from  cnscin  hv  tlio  splittiii- 
of  tJ.o  molcMilo  of  tlic  lalt.>r  into  two,  from  which  it  wonid  appear  th.v 
tho  action  of  this  cn/ymo  is  nothing  more  than  the  first  staRc  in  tl.. 
liydrolysis  of  the  casein  moh-cilc.    The  whey  nll.nmose,  ncconling  to  tlii 
view,  is  a  I)y-pro(1nef. 

There  are  many  invest i-rators.  iM.uever.  who  I.elieve  that   reiinin  an. I 
pepsin  are  not  identical,  since  an  infusion  of  the  stomach  of  a  enlf  has 
powerful  clottiiiir  action  on  milk  Imt  a  very  w(.ak  diirestivo  one  on  cl- 
wliite,  V  hereas  n  similar  infusion  from  the  stomach  of  a  piij  shows  exact K 
the  reverse  pro).erties.     This  .mestion  is  one  of  so  controversial  a   ii,, 
tnre   that    it    would    he    out    of    place    to    u-o    info    it    further   here.      I 
should  he  p,,inted  out,  hoMcvcr,  that,  \vhen  tl-c  u'astric  contents  are  a.-i.' 
in   reacti.>n,   tnilk   Mill   lM..M,me   slotted   hy   the  action   of  the  acid   its,.' 
(inite    independiiitly    of  any   |m  p>i„    „r    rennin    the   juice   may  eontai' 
This  acid  dottinjr  of  milk    i>    prohahiy   of  a   difTerent   chemical   nafn. 
from  that  prodnccl  hy  the  i  n/ynics. 

On  other  fo.Mlslufrs  fh;in  proteins  the  action  ..f  the  <.'asfrie  Juice  i 
rclativ..ly  unimportant.  ;iMrou..'h  polys;i.-cIiaridcs  may  he  consideraM 
I.rokcn  down  in  the   .ardiac  en. I  of  the  stonnn-h  on  necount  of  the  actio 

«(  s«al|,m,.,|   v;,liva     sec   pa-.'  4:.4),  and   .lisa.'charides.   as  we  have  s 

may  hccomc  split   l.y  the  hydrnlyzinir  ,.|Tcct  of  the  hydrogen   ion.     F,  • 
diu'cstion  also  t;d<(s  place  in  the  stomach  when  the   fat  is  taken   in  .t 
emidsitled  conditi..ii.  as  in   milk  and  i-j'j  yolk,  hut  not  when  in  mass,^ 
as  in  meat  or  '.utter     Tlii..  a-'tion  is  diU'  to  the  i.resence  of  a  fat-splittin 
enzyme,  or  lipase,  in  the  'jastri.'  juice. 


•tn   the  ril...\('   i;'.TtT.-'i,  I,|.,,rc  r.TKrin 


.iv  cn^riimjrm.  an.'   i.,ir.nia''rin   tli.-  >;amr   as  ra-. 


rilAl'TKIt  lAI 
THK  HIOCHK.MICAL  l'U(MKSSKS  (»l"  DICKSTION  (r..iitM> 

DIGESTION  IN  THE  INTESTINES 

'I'lu-  further  cliaiii.'cs  which  the  hnlf-<liir<'Nte<l  fmidstufTs  in  the  rhymo 
iiii(li'i({o  ill  the  iiittstiiiiil  caiiiil  tlopoiiil  oil  the  onzyinos  present  in  the 
Mcrctioii  of  Ahv  various  irlands  aiul  on  the  presence  of  Imcterin.  The 
most  important  of  the  diirestive  juices  are  the  pancreatic  juice  and  bile. 
Tlie  latter,  however,  does  not  contain  any  en/.\  iiic,  its  influence  on  di|;es- 
linn  heinjr  entirely  adjuvant. 


Pancreatic  Digrestion 

When  Hc  were  consiilei-iiiK  the  mechanism  of  se<'retion  of  the  pan- 
ri-e.-itic  juice,  we  saw  that  the  juice  jiroduced  by  the  action  of  secretin  oil 
'l;c  (iland  cells  does  not  contain  any  active  proteolytic  enzyme,  altlioni;h 
it  contains  one  cai)able  of  actinir  on  i)tilysnechnrides  and  another,  on  fat. 

TiiK  Aitkin  ok  Tkyi'siv 

Wlieii  this  juice  is  mixed  with  the  secretion  of  the  duodenum  or  of 
the  upper  part  of  the  small  intestine,  it  iniiiiediatclv  develops  powerful 
liroteoiytic  power.  The  same  result  may  also  be  obtained  by  mixinsj  it 
"illi  an  extract  of  the  mucous  membrane  of  the  iluodeniim  made  with 
'lihite  l)iearl)onnte  solution.  A  very  small  aiiiouiit  of  the  extract  is 
liipjible  of  increasinjj  the  digestive  activity  of  a  very  considerjil)lc  (|uan- 
lity  of  pancreatic  juice,  showinu  that  the  action  depends  on  the  presence 
"I'  an  enzyme  uhii'li  has  been  called  inltnikliia.'tf.  This  inlluencc  of  the 
intestinal  secretion  is  readily  destroyed  by  heatiiiir. 

I..njr(>  (juantities  of  alkali  iire  contained  in  the  pancreatic  juice  and 
I'ilc,  so  that  in  the  upper  reaches  of  the  intestine  the  acidity  of  the 
'liyme  is  practically  neutralized.  A  little  lower  down,  however,  an  acid 
I'Mctioii  may  ajrain  develop  (see  patfc  ."lO.'i).  On  account  of  these  facts  it 
ti.is  been  eonclnded  that  the  activity  of  trypsin  is  most  rapid  in  the  pres- 
•iiee  of  a  slight  excess  of  hydroxyl  ions;  i.e.,  in  a  weakly  alkaline  solu- 
lion.  It  is  interestinji  to  note  that,  as  a  result  of  the  Kreat  secretion  of 
ilkali  by  the  pancreas,  extracts  of  thi  rgan  after  death  show  a  very 
liiu'h  decree  of  acidity  in  com])ari.son  with  extracts  from  other  orjians 

4'iO 


ii'i  f 'i 

Mi' I  !    ■ 

I; 


4!I0 


DKillSTlOV 


11 


}'  .t 


!*fi 


IMl! 

niii\ 


.-iiiil  tissues,  ll  Iiiis  iils.p  recently  Iieeii  nIkiwii  tluif  tlie  activity  of  try] 
.l-.es  not  .lepeii.l  on  the  presem-e  of  free  Iiy.lioxyl  ions,  but  that  it" 
IHoeeed  in  tlie  inesenee  of  (viv  .•iei<|,  even  Up  to  a  strength  of  ("ii  ^  ].:> 
If  p<"psin  is  ]. resent  toiretlier  uitii  trypsin  iu  a  distinctly  aeid  solution 
the  pepsin  seems  to  destroy  the  trypsin,  uidess  the  mixture  contains  ,i 
consideraMe  ipiantity  of  protein,  when  the  tryptic  activity  may  persist 
even  for  several  hours.  A  i)raetie:d  conclusion  that  wo  may  draw  froi,; 
those  results  is  to  the  elTcct   ih.-it   preparations  of  trypsin-  the  so-calh-l 

IKiiiinatlii.  for  example     if  iriven  with  the   f 1,  nmy  pass  in  an  active 

'■ ''''""  '"'"  '•'"■  'Inoclennm.  where,  in  the  more  favoraMc  onvironnient 

created  l.y  the  neutrali/ation  of  the  ox.-ess  of  acid,  it  will  doveloi)  its 
proteolytic  pi,«er.  The  therajicntic  administration  of  pancroatin  i-. 
therefore,  jnstilicd  I  Loin,'"  .. 

The  actJMited  trypsin  acts  on  proteins  in  very  mucli  the  same  way  as 

pepsin,  except  that  the  d mposition  of  the  |)eptono  and  proteoses  intn 

polypeptides  is  the  chief  feature  of  the  i)roccss.  Thus,  after  tryptic 
dii:estnni  has  proccedc.l  t'or  some  time,  onlv  a  trace  of  (yrimary  proteoses 
l.ut   .•onsideralde  .|uantities  of  leucine,   tyrosine  and   other  amino  acid^ 

"ill    lie    found    present.      S ^    investii:ators   holieve    that    the   tlioronirli 

nature  of  the  di-ri'stive  action  of  activate.l  i.ancrcatic  Jnico  may  depend 
|>n  its  jilso  containiny:  erepsin.  an  cn/yiiu'  which  we  shall  sop  tr)  bo  i)rcs 

•'"•  '"  isidcrable  amount  in  the  mucous  membrane  of  tlie  intostino  and 

other  tissues,  and  whose  p:irtic\dar  function  is  to  split  i)olypeptides  into 
the    amino   aej.Is.      l-'rom    the   autolytic   digestion    wliich    takes   jilaco    in 
orirans  kept   in  a   sf,.rile  condition  after  death,  tryptic  dipostion  difTcr^ 
in  that  it  produ.-es  only  small  .piantiti.s  of  ammonia.     The  larsro  (pianti 
tics  of  ammoni.'i   pro.inccd   in  autolytic  diirestion  no  doubt  have  a   rolii 
lionship  to  the  ;icids  simultaneously  set  free  durin-:  this  process. 

Ill  tin   i,roili,tls  of  Iniiili,    iJIfitxlioii  it   is  usindly  found  that,  althou;:!' 
tliere  lijis   been   consi.lerable  splittin-r   of  the   |)rotein    into   amino  acidv 
there  are  still   ;:    (rood   many   aiiiino-carboxyl   iXIK'O')    linkajros  left   un 
I'roken,   indicatinir  that  certain  polypeptides  ar.'  left   intact   in   the  mix 

nil'.  To  split  the  polypeptides  re.piires  the  ;iid  of  the  //«;«/„.  which  i~ 
present    III    the    iiMieoiis    iiieiiibiM lie    of    the    intestine.      Interesting'    iiivc- 

tiuatioiis  have  been   ni.id the  exact   lU'iivv,'  to   whhdi   trypsin-ontei- 

kinase    can    split    up    tli.'    various    knoun    jiolypcptides.      This    seems    !• 
depend  on  the  stiuctiiie  of  ihe  polvpeplide  mol..cule  and  on  the  liumlic 

of  amino  acids   present    in    il j,;,!,,.      {'or   example,   jinalylsrlyciiie.    I,ir 

not  srlycylalanine  is  hydrolyzcd.  jiltlioiiirh  both  contain  the  same  amin 
acids  but  linke.l  toirether  in  a  different  \\a.\:  and  tetra-flypyljjlyoin. 
whicli  contains  live  Ld>ione  ra-licles.  is  liydiolyzed,  uhercas  di^'lvcyli;!- 
cine,  \\liicl nljiiiis  only  three,  is  not. 


Tin:  ii|(>(1ii;mic.\i.  ntoriissis  of  i>i(ii;sTioN 


491 


Till'  iiiiixir-tjiiicc  of  tlic  presence  of  rrrpsln  in  tlie  mucous  mcmbraiio 
n\  the  intestine  is  lliat  it  serves  iis  a  liiii-rier  to  the  passiiKc  of  any  tinsplit 
.iiiiino  aciils  from  tiie  intestinal  contents  into  the  bloo<l.  It  insures  the 
I'lcaUinK  \ip  of  tlie  pi'otein  moleeiile  into  its  ultimate  units  before  absorp- 
tion. Tlie  fuitlier  fate  of  tbe  absorbed  amino  acids  -will  be  considered 
;inder  the  subject  of  ])rolein  metabolism. 

Tin:  A(  rro.v  or  Liiwsi: 

Neutral  fat  is  decomposed  into  I'atty  acids  and  trlycerine  by  the  lipnxr 
|ii'esent  in  the  pancreatic  juice.  This  en/yme  may  also  be  extracted  from 
tlie  fflands  by  means  of  (iO  i)er  cent  ali-ohol.  Its  action  is  remarkably 
accelerated  by  the  jiresence  of  bile,  and  considt'rably  de])ressed  by  inor- 
L:aMic  salts.  It  is  also  xci-y  <lependenf  on  the  dejrree  of  alkalinity,  the 
Hptimum  beiufr  a  h\-diofren-ion  concentration  of  11x10".  The  favorinsr 
.K-tinn  of  bile  is  iniiIoubtcdl>  nwinj:  to  the  bile  salts  (si'c  pa<re  4!>:>),  and 
it  is  i)robable  that  this  action  is  dependent  upon  the  influence  which 
tliese  have  in  lowering  surface  tensioi'  and  therefore  brinsriu<r  about  a 
iimre  intimate  co'      '-t  between  fat   and  water. 


Tin:  A(Tii>\  of  Amyi.oi-sin' 

The  action  of  pancieatic  juice  (ui  carbohydrates  de])ends  <ui  the 
•nnylolytic  enzyme  called  iniiiilnpsin.  In  animals  havinpr  no  active  ptyalin 
ill  tlie  siilivii.  amy]o|)sin  serves  as  the  only  diastatie  enzyme  eoneerned 
ill  tlie  difiestive  pi'oeess.  In  any  ease,  at  least  for  the  first  stapes  of  the 
■  lisiiii)tion  of  the  starch  molecule  that  is,  its  conversion  into  dextrines — 
:iiiiylopsiii  is  a  more  powerful  enzyme  than  |)tyalin.  It  does  not  appear 
to  lie  so  eflicieiit  as  ptyalin  in  the  final  stajres  of  the  hydrolysis,  for  it 
dues  not  ])roduee  so  much  reduciitji  suirar  as  i)tyalin  does.  Indeed  ex- 
tracts of  pancreas  will  sometimes  convert  starch  into  soluble  starch  and 
•  lixtrine  with  preat  speed,  but  inoduce  scarcely  any  reducing  sugar. 
"11  this  ;iecouiit  it  is  believed  by  iiian.v  investigators  that  there  are  at  least 
two  distinct  ami  separ.'ite  enzymes  in  amyloi)siii  and  also  jierhaps  in 
|it,\aliii,  one  ji  true  aiii.vlase,  which  converts  starch  into  dextrine,  and 
I  lie  other  a  dextrinase,  which  con\-erts  dextrine  into  maltose.  In  the 
■asc  of  both  ptyalin  and  amylopsin  digestion  |)roceeds  best  in  a  very 
I  ak  acid  reaction.  Amyloj)siii,  jis  it  is  secreted  in  the  i)aiicreatic  juice, 
is  fully  activated:  bile.  ai)ail  from  the  alkali  which  it  contains,  having 
Ml  iiilluence  (ui  its  digestive  power. 

Hcsides  amylopsin  the  pancreatic  juice  also  contains  maltase,  and  in 
ill'  cast'  of  vdiiiig  aniiiiiits  or  ul'   I  hose   thai    lake   milk    with   their   food 


tiifoiighoiit   their  li\ 


;icti 


After  the  suckling  animal  has  dis- 


Pf  i  ^' 


492 


HKiKSTION 


continued  taking  milk,  the  lac-tasc  disaijpcais  from  the  pancreatic  jui. 

Attempts  have  l)een  n)ade  to  brine  it  l>aek  liy  iVedinj;  the  ndnlt  np' 

milk,  I)nt  witliout  success.  Occasionally  the  pancreatic  juice  also  en 
tains  ini-crtdsv. 


y   M 


M'^  1;^ 


^ 


The  Bile 

Associated  Avith  the  iiancreatic  .juice  in  all  its  functions  is  the  hi!' 
When  tliis  fluid  is  ju'cvented  from  entering  the  intestine,  the  digest!' 
I)rocess  heeonu's  very  inipert'c-l,  the  alis.irption  of  fat  lieinif  particiilai  J 
interfered  uith  i  see  pai;c  tilM).  I'.ile  is  alsu  an  excretory  product,  .-ii. 
its  fompositinii  therefore  is  much  mure  complex  than  that  of  the  otlp 
digestive  fluids.  This  vaiies  ver\-  much,  hnwcver,  jwcordinir  to  il 
methoil  of  collection,  liile  from  the  u'all  hhnldcr  after  death  eontaii 
much  more  solid  material,  imrticularly  hilc  salts  and  mucin,  than  tli. 
collected  from  a  fistula  of  tin-  hile  duct  (u-  trail  bladder  duriii!;  li|. 
These  difTereiH'cs  will  lie  evident   from  the  aecompanyiuir  tal)le. 

Itiic   fr..iii 

(lull   l.lM.Mor  Fi.stulii 
till)  |,;irts  .outrun     - 

\V:ili'r    Si;  i^j 

S"li'^     It  '.■? 

<)it;;uiii>  suits    il.il,'  smUs. <•  O.Ol-S 

Mucin   ;iii.t   Lil.'   |.i-t.i,'nt .1  n.'i 

rii.iiostcr.)!    (i.:2  n.od  n.ir, 

T.ccifiiiii  aii.l  f.it 0..-,  i.n  n.n2-n.nn 

Iniir^nnif    s:.|ts    n.S  0.7-0. S 

Tn  general  it  may  l)p  said  that  bile  obtained  from  a  fistula  in  m.i 
contains  only  about  H  per  cent  of  total  solids,  of  which  from  one-fouit' 
to  one-half  are  inorjranie,  whereas  bile  from  the  gall  bladder  eontaii 
10  to  20  per  cent  of  total  solids,  of  which  only  about  one-twentieth  .11 
inorganic.     The  chief  cause  for  this  difference  appears  to  be  that  wh. 
the  l)ile  goes  to  the  intestine,  a  considerable  proportion  of  its  bile  s;ii- 
is  reabsorbed  into  the  portal  bhiod  and  recxcretcd  by  tlie  liver.     S..11 
of  the  difTerence   may   also  be  caused   by  the   fact   that   absorption    . 
Avater  takes  place  from  the  gall  bladder,  and  that  mucin  and  possih 
cholesterol   are  secreted   by  this  (u-iran.     These   striking  differences  b 
tween  fistula   and   gall-bladder  bile   are   ob.served   oidy  when   the   cni 
mon  bile  duct  is  occluded.     If  the  bladder  fistula  is  made  with  the  ci.i 
mon  duct  left  open,  some  of  the  bile  trains  entry  to  the  duodenum  io 
therefore  l)ecomes  recxcretcd.     It  is  well  known  that  a  fistula  of  the  g,i 
bladder  in  man  after  a  tinse  e!os,^s  up  .iiiti  the  bile  again  takes  its  usu. 
course  along  the  I)ile  duet  into  the  dnodeinim. 


tut;  iiioiiiMMKAi.  rRoi  i.ssr.s  (if  nKiKSTinv 


493 


liitciTstiiiK  ol(scVvati()iis  liavc  Itccii  ciillcctcd  oil  the  fiinouiit  of  the  secrc- 
iii.ii  from  a  fistula  liotli  in  man  and  in  llu'  lowor  animals.  In  man  it  is 
.■,,),im<.iily  statfd  that  ah-nt  nOO  cc.  of  l.ili"  arc  secreted  daily,  the 
.iinouiif  varyiiii,'  eoiisiderahly  dnrini;  tlm  dilTereiit  hours  of  the  day.  The 
Mcretion  of  hile  is  uieatly  reduced  hy  lieni(.rrlia','e.  It  is  greater  on  a 
meat  diet  than  on  on.'  of  carhohydrates.  It  is  reduced  during  stara- 
iioii,  hut  eoiitinucs  to  he  secreted  up  to  the  moment  of  death. 

Fl  NCTKIN'S  OF   IJlI.E 

One  of  the  main  functions  of  the  hile  salts  is  that  they  greatly  assist, 
not  only  in  the  dii-'estion,  hut  also  in  the  ahsorption  of  fats.  When  bile 
IS  excluded  from  tlie  intestine,  the  feces  are  loaded  with  fatty  acids 
^^llich  have  lieen  split  off  partly  hy  the  now  less  effective  lipase  and 
,,artly  by  the  action  of  hacteria.  The  fatty  acid  thus  liberated  in  the 
iihseiiee  of  hile  salts  is  not  absorbed,  because  the  bile  salts  serve  as  the 
rarriers  of  fatly  acids  into  the  epithelial  cells  and  laeteals.  They  com- 
l.ine  with  the  fatty  acids,  probably  by  formintr  some  chemical  compounds, 
ill  which  they  carry  th.'in  into  the  endothelial  cells  where  the  compounds 

I nie  disrupted,  the  fatty  acid  ccunbininir  with  glycerine  to  again  form 

iM  utral  fat  and  the  bile  salts  being  carried  to  the  liver  and  reexcreted. 
Tlie  innuence  of  bile  salts  in  assisting  the  action  of  lipase  is  probably 
,lue  '■!  a  lowering  of  the  surface  tension,  thus  bringing  water  and  fat 
into  closer  union.  This  accelerating  iiitluence  has  also  been  demonstrated 
•shell  sviithetic  bile  salts  have  been  iisc.l,  sh..wiiitr  clearly  that  it  is  really 
ilicse  and  not  any  other  constituent  of  the  hile  that  are  responsible  for 
It-,  accelerating  intluenee.  » 

Uilc  also  functionates  as  a  rcgiilatur  of  inlfstinni  pntrrfaction.  This 
It  does  apparently  because  of  its  slight  laxative  i)roperties,  by  which 
the  intestinal  contents  are  csixllcd  before  the  bacteria  have  grown  to 
ativ  sreat  extent  in  them.     I'.ilc  itself  is  a  favorabh-  <'utture  medium  for 

■  rtain  bacteria,  so  that  it  can  have  no  antiseptic  action.  Its  assistance 
in  the  action  of  t-ypsin  and  amylopsin  depends  very  largely  upon  the 
iilkali  which  it  contains. 

.\s  an  fxcrcioru  veliiclc  bile  is  important,  because  it  possesses  the 
power  of  dissolving  cholesterol.  Toxins  and  metallic  j.oisons  of  various 
kinds  are  also  excreted  in  it. 

Although  not  directly  concerned  with  the  digestive  function,  it  will  be 
■  oiiveiiient  to  say  something  hero  concernitig  the  chemical  unturc  .^nd 


■rival ion  of  the  vari 


ous  biliarv  .  .mstituents. 


i  .  m      ' 


f  !t 


Hi 


'         i*..||: 


494  nrdisTioN' 

THE  CHEMISTRY  OF  BILE 

The  Bile  Salts 

Til  'iinsf  aiiiinnls  tlif  Iiilc  salts  consist  (it"  tin-  smliuin  salts  of  tflypoclioli.' 
and  taiirorlioli.-  acids.  Kacli  of  these  acids  is  composed  of  a  part  callr.l 
cholic  acid  wliidi  is  more  or  less  related  to  cholesterol,  and  of  t'lycii,, 
(t"H_.\U,C(»(>H  ainino-aceti"  acid)  or  taurine  I  ( '.HA'SO.I,  a  derivatix. 
of  e.vsteine,  which  is  u-.,miMo-/i-tliioi)ropionie  acid  (("II,HS.CII\II 
COOHi.  The  exact  form  of  ciiolie  acid  varies  in  different  animals,  Ili.it 
of  the  piir,  for  erample,  heinir  different  from  that  of  man.  Bile  salts  aiv 
nn  exclnsi\e  pr-odiict  of  liver  melaliolism:  i.e.,  they  are  not  formed  i' 
any  other  i)art  of  the  animal  hody.  Tluy  uive  a  very  sensitive  coli.r 
reaction  known  as  Pettcnkofer's.  which  however  is  not  sjiecific  of  liile  acidv 
siin-e  it  is  also  <.'iven  liy  olei<'  acid  and  li.v  many  aromatic  snlistani'cs  i\w\ 
alcohols.  It  must  he  rciiu'mlteifd  tin*  the  part  of  the  hile  salts  that  is 
ciniracteristie  of  the  |i-.r  is  the  clioIic  acid,  the  taurine  and  irl.vcin. 
lieinjr  i)res(>nf  in  other  lissiu's  and  orjians. 

When  choiic  acid  is  <riveii  to  animals  mixed  witli  the  food,  the  amoui.t 
of  taurocholic  acid  excreted  with  the  hile  is  increased,  indicatinsr  Ih.i; 
there  must  he  a  .store  of  taurine  avnilahle  in  the  orjianism.  This  sto', 
can  not,  however,  he  larire,  for  if  the  feedini;  with  cholic  aeid  is  reiteatcl 
several  times,  it  will  l>e  found  that  the  taurocholic  aeid  diminishes  an.! 
>rly<M)cholic  iieid  takes  its  ])lace:  and  this  inereased  excretion  of  ".'lye 
cholie  aeid  <;oes  on  .just  as  lonj;  as  cholic  aci<l  is  fed.  The  reserve  •>'' 
taurine  in  tlic  animal  luMly  ;ippcais  thcrefoic  to  he  limited,  althou-rh  it  i^ 
used  in  preference  to  trly<'ine  when  there  is  an  excess  of  eholie  aeid  to  1..' 
neutralized.  On  the  other  hand,  tlie  store  of  <rlycitie  seems  to  he  inexliaus' 
ilile.  That  there  is  no  resiTve  (•''  cholic  ai-id  itself  in  the  liody  is  indicated  1 ;. 
the  fact  that  no  increase  in  taurocholic  acid  excretion  hy  the  bile  resuh- 

when  cystine,   the  mother  substance  of  taurine,   is  ^'iven   with    the   f 1. 

If  both  taniine  and  cholic  acid  be  ffd,  howevci-,  the  e\<'rction  of  taiir.- 
eliolic  acid  increases. 

The  reliitive  amounts  of  taui-ocholic  and  ylycocholic  acids  in  the  bile  ^i' 

different  animals  differ  considerably.       Human   bile  itains  relati\i;\ 

a  small  amount  of  taui'ocholic  ai'iil;  on  the  other  li.-ind,  ihe  bile  of  the  d.i  - 
contains  a  larjje  excess  of  it. 


Cholesterol 

In  hnnniM  bile  the  i)ercei:tasre  of  this  importan'  substance  is  not  liii 
(  l.(i  parts  per  lOOo.,  i,ut  it  is  of  -rreat  clniical  importance  because  of  ll 
fact  that   it   nniy  sei>arate  out  as  a  precipitate   formin;;  t/dllsloncs.     Tl 


Tin;  iiKH'iiKMif'Ai.  rK()(i;ssi:s  up  hkiistiov 


49.') 


|MTcoiita','o  of  clioli'storol  in  tlifsc  varies  from  '20  to  !•() ;  tlio  remniiulor 
lMiii<r  orpaiiie  material  sueii  as  epitlielial  eells,  iiMirjraiiie  salts,  pitrmeiit, 
ttc.  The  origin  of  cholesterol  is  partly  ondofienoiis  and  partly  exof?e- 
iKiiis.  In  the  former  ease  it  comes  from  the  envelope  of  red  blood  eor- 
imseles  anil  from  the  nervous  tissues,  where  it  is  present  in  eonsidcrable 
;iiiiount.  The  latter  sonree  is,  of  eofirse,  the  food.  The  increase  in 
cliiilesterol  esters  in  the  Mood  after  feedinj;  with  food  rich  in  this  sub- 
sl.iiiee  has  been  shown,  parlieidarly   in   rabbits. 

That  the  bile  should  be  the  pathway  tliron!.'h  whieh  eholesterol  is 
excreted  depends  no  doubt  on  the  fa'-t  that  it  contains  bile  salts,  whieh 
iilciMf;  with  their  other  proi)erties  have  a  remarkable  solvent  action  on 
liiolestero  .  This  s.ilveiit  (iroperty  depends  on  the  eholie  acid  part  of 
tlic  bile  salts,  which,  as  already  remarkeil,  is  chenrn-ally  very  closely 
ivlated  to  cholesterol:  indeed,  the  relationship  is  so  close  that  some  have 
Mi^rs-'ested  that  cholic  aciil  is  derived  from  cholesterol.  This  would  mean 
that  the  cholesterol  of  blood  is  excreted  in  two  ways,  as  eholesterol  a. id 
,is  eholie  acid.  Other  observers,  however  maintain  that  the  cholesterol 
is  cNcreted  mainly  by  the  linins:  membrane  of  the  s-'all  bladder,  and 
tliiit  this  oxi)lains  why  trall-bladdcr  bile  contains  more  of  it  than  (is- 
tula  bile.  This  evidence  is,  howevi-r.  not  v  ry  stronir,  for  the  <rfPater 
excretion  of  cholesterol  under  conililions  where  the  circu'  'ticui  of  bile 
is  L'oins:  on  may  be  explained  as  due  to  the  i»resence  of  bile  salts,  wliieh 
^eive  to  carry  the  eholesterol  out  of  the  blood. 

Many  problems  remain  to  be  elucidated  in  connection  witli  the  metabolic 
History  of  cholesterol.     That  some  of  it  is  absorbed  when  cholesterol  is 

niitained  in  the  food  mijrht  seem  to  indicate  that  its  soun-e  is  entirely 
I  xi>m'nous.  Airainst  this  view,  however,  stand  two  facts:  (11  that  the 
.  Iiolesterol  in  the  feces  of  herbivorous  animals  is  of  the  same  variety  as 
tli.it  present  in  those  of  carnivorous  animals  and  not  the  i>hytoster(d 
vliich    is   present    in    plants;   and    t'l)    that    the    miivcisal    iiresenee    of 

liiilesterol  in  cells  indicates  that   it   must   be  manufactured  there. 


'     i 


The  Bile  Pigments 

The  iiijtments  of  bile  are  liilinihin  and   hllirinliii.     The  latter  is  pro- 
■liieed    from   the  former  by   oxidation.      If  the   oxiilation    be   carried   a 
=^  -:iitrc  further,  a  blue  pinmenl  cai!e<l  bilicyjinin  is  I'ormed.     This  process 

'  t'  oxidation  can  be  observed  in  the  rin;.'  test  for  bile  pigment  with 
Miiiiiiis;  nitric  aci<l.  When  bilirubin  is  reduced,  urobilin,  one  of  the 
;>'i;ments  in  urine,  is  formed.  Hilirubin  must  therefore  be  considered 
s  the  mother  substance  of  all  these  pigments,  and  it  is  of  interest  iit 
'  "iHicction    with    its   derivation    to   know    that    it    has   the   same    foiinula 


496 


f>uii;sTio\ 


^  rJ 


i# 


t   !; 


■.Ml 


:■; ! 


i:'' 


as  iroii-fri't'  luMiiatiii  or  liPiiiafoporpliyriii,  which  is  proclueod  by  treiitiir.' 
hemoifh)l)in  with  eiiiioeiitratcd  sulphuric  acid. 

Chemical  invcstipation  has  shown  that  bilirubin  is  built  up  from  suli 
stitutod  pyrrols,  jirobably  four  such  bcin^  contained   in  the  molcciilr 
The  pyrrol  Rrou))  is  also  present   in  indole  and  tryi)tophane,  and  cmi 
sists  of  four  carbon  atoms  and  an  Nil  ^roup  linked  tofjether  as  a  riii'.' 
I  set"  pajre  (!()4).     Similar  pyrrol  derivatives  can  be  i)ro<luced  by  deeoni 
posinj;  ehlorophyl,  the  jjrcen  colorinjf  matter  of  ))lants.     It  is  important 
to  remember  that  bilirubin  is  acid  in  nature,  and,   therefore,  can  com 
bine  with  alkalies  to  form  salts.     The  relative  amounts  of  bilirubin  anil 
biliverdin  vary  in  the  bile  of  different  animals. 

When  these  piKments  enter  the  intestine  they  arc  reduced  to  urobilin, 
part  of  which  passes  out  with  the  feces,  another  jtart  beiuf;  absorbed  into 
the  blood  and  excreted  in  the  urine.    Part  of  that  excretctl  in  the  iivini 
exists,   however,   as   a    so-called   chromo(>;eii    named    urohilino'ini.     Tli'' 
urobilino<jen  is  converted  into  urobilin  by  the  action  of  oxy<icn. 

The  method  h]i  whieh  nrohilin  ix  profhieed  from  blood  pigment  liiis 
been  studied  by  histologic  examination  of  the  liver  i)articularly  of  biiil-- 
and  amphibia,  in  which  destruction  of  blood  pif?ment  srocs  on  rapidly 
Increased  destruction  of  blood  pigment  can  be  induced  by  poisoiiiui: 
with  certain  substances  such  as  arseniureted  liydro<;en.  From  sucli 
studies  it  is  usually  believed  that  the  bile  piiiments  are  a  peculiar  product 
of  hepatic  activity,  beinji  ])roduced  from  blocxl  i)i'fments  that  are  dc 
rived  from  erythrocytes  which  have  been  britken  down  either  in  the  livn 
ilNclf  or  in  some  other  vi.>icus  (e.  <:.,  the  spleen).  Whipple  and  Ilooiui 
have  brouRht  forward  sceminsjly  incontrovertible  evidence  aprainst  mu-h 
a  view.  They  have  found,  for  example,  that  the  bile  pigments  an 
formed  just  as  readily  in  animals  in  which  the  circulation  of  the  livci 
was  greatly  curtailed  by  anastomosing  the  portal  vein  with  the  Yeii;i 
cava  (Eck  fistula)  as  in  normal  animals.  Even  when  the  circulation 
was  limited  to  the  anterior  end  of  the  animal  (head  and  thorax)  bit' 
pigment  a,  .)eared  in  the  blood  when  heraolyzed  erythrocytes  were  in 
jected,  and  it  was  also  formed  when  hemoglobin  was  placed  in  the  plcuiji 
and  peiitoneal  cavities.  The  endothelial  cells  of  the  blood  vessels  aii'i 
elsewhere  can  evidently  form  the  pigments,  at  least  when  the  liver  i- 
absent.  When  such  a  process  occurs  under  normal  conditions,  it  is  quit' 
probable  that  the  liver  acts  merely  as  an  excretory  organ  for  the  p\'2 
mcnts  in  the  same  way  as  the  kidney  docs  for  urea.  Possessed  of  end" 
tlielial  cells,  the  liver  might  itself  also  produce  some  of  the  pigment- 
l)Ut  no  more  than  other  orcrans  with  a  similar  number  of  those  cells. 

Even  the  derivation  of  bile  pigments  from  hemoglobin  is  called  i: 
question,  for  the  same  workers  have  observed  that,  whereas  the  excn 


TUT.    IlIOClir.MIrAt.    njlHlSSIS    (iK    DKil  STIIiV 


407 


til 


,1 ,r  i.i._M.i.Mi1   tVoin  i.  I.iliiity   listul;.   is  iTinnrUi.l.ly   ruu-<\:w\   i I'x.' 

,,,|    „„    J,    tixr.l    niixi'.l    .li.'t.    i1    I lun.-    ilMT.'iisr.l.    m.i.icI  ilil.'s    l,y     |(MI    p.T 

,.,,,1,  ulini  111-  di.'t   vv.-is  rlwili!.'.'.!  I..  "I r  .•.•nl".l,MlrMt.s.  ami  .Irprrss.-l 

,,„  ;,  ,|iH  of  ll.nil.  Til."  qlMSlinll  alis.'S  iis  tn  vv  lu't  ImT,  .ifl.T  ;ill,  tl,..  I.ll.' 
|,i..r|,„.|its  iiiv  ivally  .l.Tivfil  In.in  l.n.k.'n-ilowi.  lM'ni.iL'l..l'iM,  Mny  tln-y 
„nt    lie    niiiimfiicttirfd    '/'     "'"'"    ""'    "''   "''"'''    ii>ii'<i'i'i'^  •' 

Wliippl.'  aiHl  IIn„p<T  l.av.-  also  sliowii  tti.-.t  l.il.-  is  a  most  impottinit 
s.vn'tion.  for  .lo-s  niivly  surviv.-  or.  an  onliiiary  .li.t  if  l.il.-  is  p.-niia- 
M.iitly  pr..v.'ii1.Ml  fn.iu  .•nt.-nii!.'  tli.'  int.'sti....  Int.stinal  Nyt.ii.t..iMs 
.nun  "snp.Tvn.-.  aii.l  l..'.M.in.-  pro-.'ivssivly  iii..r.'  vv.'iv  until  tli.-  .l.-ath 
,,r  llio  animal.  FcMJins;  witli  l>il.'  .lo.'s  n-.t  irli.'v.'  tlio  .•oncliti..n,  Imt 
f.MMlintr  witli  .•ookc.l  liv.T  s.m.mis  t..  have  a  iM'ii.'ti.-ial  .'tV.-i't. 

Aft.T  .'xtinviisation  I'f  l'l.»'.l  i"  tli.'  snl.i'Ulaii.ons  tissu.'s,  as  in  a  lifuist-, 

for  .■xamiil.'.   n   .l.M-..mpositi..n    .>f   In'mo-loliin    \>v«,- Is   .piit.'    liU.'    that 

nrruii-insi  in  til.'  liver,  an.l   l.-ii.ls  to  tii.'  pro.lu.-tion   «<  \,\nv  an.l   Inoun 

.■,i„l  ..'r.vn  ).i-m.-nts  lik.-  Ili.is.-  of  tlic  l.il.'.     Wlnn  lHnio|ysi>  iv  |.io,lii 1, 

;i.  l,y  iiilialation  of  ars.'ni\ir.'t.'.l  liy.ln.tr.'U  .-r  tlu'  inj.ction  ..f  iiiorviani.' 
,,!•  KinloL'i.'al  licmolysins,  tli.'ic  is  an  i.iini.diat.'  iiicr.'asr  in  tli.'  aiiioUHt 
of  l.il,.  pitrm.'iit  in  tli.'  l.il.'.  Kv.n  th.'  inj.'.-lion  '.f  li.'iiio-lol.in  •solutions 
li.Ms  this  .ir.'i-t.  rii.l.T  th.'s.'  .•onilitions  of  li.'molvsis.  h.'sid.'s  an  in.-r.'as.' 
in  ni'..l.ilii!,  tli.T.'  may  1..'  .'..nsi.l.Tal.l.'  .piantit  i.s  of  h.i..o..rlol.in  M'.'r.'t.'.l 
ill  the  iirini'. 

Hil.'    salts   an.l    piymcnts    usually    a mipany    .a.-h    ..tliir    when    nny- 

iliiinr  o.'.'urs  to  int.'if.'i-.'  with  tli.'  frc.'  s.ciction  of  hil.'.  For  .'sampl"'. 
nt.'r  liuatioii  ..f  th.'  l.il.'  .lu.-t  l.oth  l.il.'  i.i'-nn.'iils  an.l  l.il.'  Milts  a.'.'iimu- 

hit.'  in  the  I.I0...I,  in  the  serum  of  whi.-li  ih.'y  may  I..'  r. -iii/.e.l  l.y  the 

..r.linary  ehemieal  tests  in  fr..m  f..ur  t..  six  h..urs  aft.'r  th.'  op.Tation. 
If  the  a.'eumulath.n  he  allowe.l  t..  i.n.r.'.'.l  fnrth.r.  the  l.ile  pivm.nts 
iM.n.me  .lep..sit.'.l  in  the  tissu.'s.  v'lvin!:  th.'ui  th.'  i..'.'uliar  y.llow  ish  a].- 
l-iaraneo  known  as  .jaun.li.'.'.  rn.'.er  tli.'s.'  .■..lutitioiis  th.'  hil.'  salts  an.l 
l.i'.Miients  also  app.'Jir  in  the  urine.  The  ae.-uimilatioii  of  l.ih'  salts  in 
111.'  h.i.ly  affeets  .•.•rtain  jthysiolosri.'  pr.M'.'ss.'s;  for  ..ii.'  thiii'_'.  it  eauscs 
::  '-'reat  leii<rthenin<r  in  the  elottiny;  tiiii.'  of  th.'  Mood. 

If  the  1.1. .0.1  supply  f.)  the  liv.'r  is  interinpt.'.l  hy  li-ation  ..f  the  p..rtal 
,.iM  an.l  hepatie  artery  at  the  same  time  that  the  hi!.'  .liu'ts  are  o,...hi.l.'.l. 
n.it  a  tra.'e  either  ..f  l.ile  salts  or  of  l.il.'  pi'-'m.'nt  api..'ars  in  lh.'  l.h.o.l 
.buiii.;  the  six  t..  eii:hteen  h..urs  tluit  th.'  aiiimaiN  Mirviv.'  tin'  ..p.'rath.n 
The  am..unt  of  ..l.struetion  ..f  the  l.ile  .In.'t  n.'.'.'ssary  to  j.ro.lil.'e  these 
symi.t..ms  is  very  slitrht,  siii.-e  l.ile  is  s.'er.'t.'d  at  a  v.'iy  low  pr.'ssure. 
<-!ol   of  liiueus  or  a  swolUii  <•..!!. lili..n   "f  the   mii.'..iis  111. ml. ran.' 


r.-.  rn 


if  the  .luct  is  sufliei.'iit  to  i.r...lu.'e  ohstruethm.     In  the  .lis.-liari:.'  of  hil 
from  the  iiM  hliuWer  into  the  .lu...leiiiim  it  is  elaii I  l.y  .M.'lt/.'i'  tini 


4 

i! 


t  a 


4»8 


I>l()KSTIO\ 


•il 


iiM'iinocal  relationship  exists  iM'twicii  tin-  (Miiitinction  of  tlio  lilmM. 
musculature  ami  the  relaxation  of  the  muscular  fihers  surrouiulinK  Ih 
tluet  iu  the  duodenum.  If  this  reciprocal  innervation  fails  to  operai 
properly,  discharge  of  bile  into  the  duodenum  may  liecome  obstruct) 
so  that  a  certain  amount  passes  back  into  the  blood,  as  in  cases  of  bil< 
duct  obstruction. 

Mile  also  contains  a  certain  amount  of  In  ithlii  and  other  phospholiiiin- 
The  amoinit  varies  considerably  in  the  bile  of  difTerent  animals,  even  it 
animals  of  the  .same  species.  It  is  probably  derived,  as  already  men 
tioned,  like  the  cholesterol,  from  the  breakinR-down  of  red  LKhmI  eoi 
puscles  that  ffoes  on  in  the  liver.  It  is  no  doubt  diRi'sted  by  the  ferment 
of  the  intestinal  tract,  the  liberated  cholin,  siiu-e  it  is  toxie  if  absorbc.l 
bcinjr  further  attc.cked  by  bai-tcria  so  tis  to  become  converted  into  cii 
tain  substances  of  a  nontoxic  luiture. 


i  ! 


im ' 


r   t 


\ 


i  -i 


CIIAI'TKU  lAII 

HACTKKI.M,   DUiKSTIdX    IN   TIIK   INTKSTINK 

On  ill)  !iv»riii;('  ilit't,  in  twcnty-fniir  lionis  the  ff<'<'s  id'  nnm  wcijfli 
aliont  KM)  iriniiis,  or  ai'ttT  .Iiyini:.  iilxnit  '20  siiaiiis.  Alu.ut  oiic-fouilli  of 
till'  (li-v  iiiiittcr  consists  of  tlit"  )>o<lics  of  liactcria.  If  i)latO(l  out  liy  tlio 
nidinary  liactcrioloirii-  ni.'tliods.  how f\ cr,  it  will  \>v  foun.l  flint  only  n 
small  propoilion  of  liirs.'  Iiactnia  an-  livini.'.  Tin-  ^'i-catcr  numlicr  have 
lici-n  (lostroyctl.  [iroliaMy  t>y  tlic  action  of  tlio  inuciii  in  tlif  larjro  intes- 
tine. Tlie  nitrogen  content  of  tlie  feces  amounts  to  aliout  1..')  <iiams  a 
ilay,  of  whieli  abont  one-lialf  is  haeteiial  nitroiren.  If  the  diet  eontains 
larse  (luantities  of  cellulose  material,  as  in  irreen  veiretalile  food  and 
I'lnit,  the  mass  of  feces  as  well  as  the  bacterial  c.nitent  may  he  consid- 
I  fahly  tri'oatei-. 

The  foi-eiroinu'  ftiets  indicate  that  very  extensive  l.aeteriolo<;ie  proo- 
cnsos  must  he  ^oinir  on  all  the  time  in  the  intestinal  contents,  and  the 
.piestion  arises  as  to  whether  siK'li  action  is  heneficial  oe  ..therwiso  tn  the 
.niimal  economy.  To  answer  this  (|uestiou  interestinir  ohservations  have 
l.een  made  on  the  <:rowth  and  well-l.ein<r  of  animals  excised  from  the 
uterus  iiuder  strictly  sterile  conditions  and  maintained  thereafter  on 
sterile  food.  Such  ohservations  made  on  <iuinea  liijis  have  shown  that 
the  animals  thrive  and  jrrow  perfeetly  for  a  considerahle  time.  Experi- 
ments carried  out  un  chicks  have  not,  however,  yielded  similar  results, 
'hicks  hatched  out  from  the  ("S<Z  under  strictly  sterile  conditions  and 
ihen  fed  on  sterile  Rrain,  do  not  thrive,  Imt  do  so  if  with  the  tfrain  is 
mixed  ji  certain  amount  of  fowl  excrement.  These  experiments,  appar- 
ently eontnidictory  in  their  results,  .show  that  for  certain  !.'rou))s  of 
iminials  hacteria  are  re<|uired,  hut  not  for  others. 

Tlie  difference  is  prohaMy  dei>endcnt  on  the  nature  of  the  foods.  It 
uill  he  rememhered  that  the  size  of  the  lar-re  intestine  varies  consider- 
.il.ly  accordinir  to  the  nature  of  the  diet  (see  pa^'c  Hi:'.).  Animals  takinc 
u'reat  (lunntities  of  celhilose  foodstuffs  have  very  larjie  ceca  and  very 
lonir  lar>,'e  intestines:  whereas  those  which,  like  the  cat,  live  practically 
entirely  on  cellulo.se-free  foo.l.  have  a  rudinientary  larfic  intestine.  The 
si/.e  of  the  lower  intestine  is  ohviously  dependent  on  the  presence  or 
.;!.s.iice  of  cellulose  in  the  food.  It  will  he  rememhered  also  that  the 
forward  movement  o      ' 


I! 


!     .'i; 


indeed,  special  provision 


is  made,  hy  the  presence  of  the  so-called   anti- 


4»9 


5  I 

!. 


■t 


H 


:t,ii 


jftii 


n  '' 


'■^'     ' 


.ilNI 


IHIIISTIOV 


iMiisliilli.'   \\.i\r,  to  (Irliiy   its  li|<>\  inii'llt       This  silU't't^ts  llmt    nil   illl|Hirt||lit 

''i-'"''i^''  I"' ■"■>  '"iiisi   Im-  iM.iriTilini.'  ill  this  jiiirt   i.i'  thr  nut       In   this. 

Uil.vs    i'<iii.|ilinii>    lirmiiii.    ,sl;il.||s||,'.l    in    t|ir    ri'iMllil    fill'    the    ili'liM-    .i|(ci;i 

lien  ..f  hii.t.iiii.     ■rii.v   iittii.  k   ll ■Iliih.s.v  nil. I    lih.Tiitv   th.-  iiioiv  .li^n.^ 

lil.lf  lu.i.lsliiirN  <'i>iitiiilii'.l  ill  thr  Nfu'clillilf  .•ells.  iiU.i  prmlliciiiir  ,,||t  ,,1 
th.'    .rlllllMsr     its,.|r    |||J||,.|ii,|>    ,,r    iMllllliv.-    Villll.-.        Thr    ilri.ls    Ihlll     ilfv    ills,, 

I"'"'' I     '■>     "li-*     |.l"c.ss    HIT     liilllliili/,.,1     |,y     thr     riil'lioiiiitrs    scrnli,! 

Ii>    the   lllllriisii. 

Ill     .■.iliiiii     h.ll.i\..rni|s    illlilllills      liif     llliiiillMlits      tliis     pniccss     ill     thi' 

•  ■(■rlilii    is   11. . I    r.l,ili\,.|y  .,!•  sll.'h    illl|>.>l't:i Ii...';nis..   it    1;ik,.s   p],,,.,.   j,,    1 1,,. 

I'.iiiiirli.  Th.'  .•iiiiniiils  s\will..\v  thi'  f.Mi.l  .111.1  it  mixes  in  this  pnrt  ..f  th. 
st.iiiiii.-h  uitli  Ww  s.-ilivn.  s.i  th.ii  hii.M.'iiii  .•iii.l  tViiii.nts  <Miiit:iiii<'.l  in  it, 
.-.■ill. '.I  ,v  iMs.'s,  jittii.-k  Ih.'  .•.•Ihiliisc,  lih,T;itii|..'  the  in. .re  cjisily  dincslcl 
I'.HMlstiiils  ii,,l..s.,.|  within  th.'  .-.■11  Willis,  As  this  pr.„.,-ss  p„^s  on  iici.U 
.■i.-..iniiiiliit.'  in  til.'  .liu.'siix.'  nii\tnr.'.  'I'h,.  f.,..,!  is  then  rctunicl  t.i  th.' 
III. .mil.  .'h.'U.'.l  .,\,.r  .'iiriiiii,  iiii.l  suiilh.w.'.i  imiiin  int..  the  niiiin  st.nu.i.'h. 
wUrvr  it   is  .li._M.st.'.l.     Th,.  iii.l  whi.'h  hii.-t.'i'iii  r.'ii.hr  t.>  .li).'.'sii.,„  .|,.p,.n.|s 

""'"''"' ''"•    "il'll"'    <>(    lllf    lii.'t.        .Mill,.     Il.'il|.,r    .,nilli\.,l.>llS,    stllM.ls     llli.l 

Uiiy  Ipi'lw.cn  th.'  t\\..  m'..iips  ..f  jiniiiiiils  .jis.'iiss.'.l  nlM.vf.      AlthnllKh  tli.' 

'■•'""I"""'  '-'"itiii I  in  his  r,„,.|  is  lint   iis.'lf  siii1i,'i,.|,tly  .Iijr..st.'.l  to  I'liniisli 

nutiiiii.nt,  y.'t  it  is  s..  li,,  ;i,.|,.,l  up,,||  ,1^  t..  p.'iinit  tin-  riiptiii'.'  ..f  th.' 
.•I'll,  th.'  .•..nlcnls  ,,f  whi.'h  .'ir,'  ih.i,  .|i-.'st,.,|.     Th.'  .•.'Ilul..s.-  is.  li..w.'v.'i'. 

•  >(  vjilii.'  ill  tiiiiiisliinir  hulk  t.>  th..  iiit.'stiiiiil  .•..iit.nts  '•intcstinn!  luil- 
iiist."  it   is  siiiii..tiiii..s  .'iiilc.l. 

Ill  til.'  siiiiill  int. 'Still.'  in  Miiiii  tJK'i'..  ill',.  Iiii.-t.'ii.'i  .•iipiil.lf  ..r  ficiiiijr  ,,i, 
••:iil...h,MliJit«'s  iiii.j  pi'.i.|ii..iii..'  ir..iii  thciii  ..I'ljiini..  .-i.-iiis,  su.-ji  ns  lii..ti.'. 
ii.'.ti.-.  It.'.  S,.  I., lis;  iis  11  slll1i.M..n.'y  ..f  .•;ii'l..,|iy.|i':it|.  exists  t.i  eneniirim.. 
the  ii.'ti..ii  ..f  tliese  l.ii.-tt.i'iii.  ..ili.'is  hiivini.'  .Mn  ii.-tmii  .>n  pn.tein  do  ii.,t 
seem  t.i  thrive.  It  Ituiy  he  flnil  this  is  t.i  he  il..e..iiiite.l  t'.ii'  piiitly  1)\  tli. 
lini.lil.'tiiin.  .It'   ii.-i.l    siihstiin.'.s    by    th.'    .•ailM.hv.ii'Mte    1.  -  ii!eiitiiti..n,    iiii.J 

pilltly     l.y    the    lii.-t     thiil.    iis    s i    iis    the    prnteiii    in. .1. '.-ill.,    is    l.i'.ik.'ii 

.l.iwn  liy  the  .lii:..stive  eii/ymes,  its  hiiii.liiiir-st.ine  iiiiiiiin  iieids  are  ;ili 
s..i'lie.I.  Tht'i'.'  ill-.'  pi'.ihiihly  ills,,  lijieteiiii  in  the  small  intestine  eiipalil.' 
nf  siilittiiii.'  fill  int..  fatt\  ii.'i.j  an. I  (.'iyeerine,  l.iit  prii..ti..ally  n.ithiiijr  i- 
kii.iun  ..f  tli.'ir  ii.'ti.in.  In  th..  liirj.'..  int.'stin.'  .if  mini,  al.m;:  with  tin 
eelllll.>s,.--lii.'-(.stiii«  liii.'teiiii  iili'eii.ly  nielit  i.ilie.i.  pr.)t.'iil->li!:est  iliii  Iiiie- 
teiia  are  ;ils..  pres.nt.  Tlies..  I,;iet..ria  l„.|,,iijr  t.t  the  eliiss,  I'.a.'illus  enli 
e.iiiiiiinnis.  th.'  Villi. (lis  iii,'iiili..|s  .if  whi.-h  are  kiinwn  iis  fii.'uhitiv.'  ;ina. 
r.>l„'s  li.'.'jins.'  ihey  .-ail  !.'r.>\\    in   lli.'  presence  .ir  alis.'nee  .if  .ixyireii. 

If  l.ii.-t.'i'ial  •rr.iwtli  is  .'x.'i'ssiv,'  ..r  tlier.'  is  an  insunieieney  .if  eiirl>..liy 
.iiiit.'s  in  til.'  small  intestine,  the  lia.-teriii  iittaek  th..  iimiii.,  aeiiU  iir,, 
.itieed  l>y  the  disrestive  eii/yiiies  an. I  dee.imp.ise  tli.-m  info  proiiuit-. 
tliat  iiiiiy  Im'  toxic  if  ;il)s.>rl).'.l  info  flie  Moo.l. 


iiA(  Ti  Ki.vr.  i.nii;sTi(i\  IN  Tin:  intoiim; 
Bacterial  Digestion  of  Protein 


:m 


I'l'iiin  .'I  |ijitliiil<iuic  stiiM<l|ii>iiit,  till'  must  iiii|«irliiiit  iH'tinn  nf  luM'tiM-iii 
i>  tliiit  wliirli  liikcs  |il:ii-i'  nil  |iii>li'iii.  rinlri  iiiiMi'i nliji-  <-<iiiclil  iiiiis  the 
iiitiNtiiiiil  liiictcfiii  liii\('  ill  '.'riit'iiil  Itif  piiwcr  id'  siililtiiii:  "IT  llir  iiiiiiini 
'.'lolip     \\licltil>     llinlcr    iiclipliic     ciiililil  inns     llirv      split     iilT     till'     i';irliux\l 

'.'idiip.  This  splitliiii;  otT  iil'  tin-  rjnlmwl  ttniiip  as  i-.ii'linii  ilinNiilr  is  pn- 
lii;iili-i|  li.v  llii' sii  r;illi'<l  i'.illiii^>  Ijisi'  liiirtrlia,  iillij  it  iiiji.v  tllkr  plarr  iitinr 
licriiTi'  ""I'  al'trr  ili'aiiiiili  timi  ( sii'  pair''  til'n.  ll'  it  liappi  lis  al'tff  this 
priii'i'ss.  thr  prmlllrts  aii'  lint  liiuhlv  tii\ic  ami  iiifliiili'  phi'iiol,  i-ri-snl, 
iinliili'  jiiiil  skatiili',  whirh  aii'  piiitiy  alisnilii'il  into  the  lilmiil  ami  partl\ 

CMMl'fl'd    W  itll    till'    fl'fl'S.    . 

Till'    friictioiis   lit'   thosi'   siilistai s   that    ari'   alisoiln'il    iiiln    thr    IiIimmI 

h.'iM'  tlii'ir  tiixii'itv  i'i'iiii>\  ril  liv  I'liii.jiiijatiiiii  mainly  with  siilphiirii'  iii'i'l 
111  I'lii'in  till'  so-i'alU'il  illiiiiiil  siilitlniti  s.  A  part  is  also  rnmliini'il  with 
•/l.M'iirnnir  ai'iil  (sit  paiii'  ><\-^.  In  tin'  i-asi'  nf  phiiK.I  ainl  m'snl  this 
iniijiiKiititiii  iH'i'iirs  imiiH'iliati'ly  al'lrr  alisnrptinii,  luit  in  tin'  I'lisf  nl' 
iiiiliilc  aini  sixafiilr  it  is  pri'mlril  liv  an  oxiilatiM-  prnrrss.  n,ii\  ritiiiv' 
ihi'si'  siilistanri's  iiitii  iinlnxvl  ami  sUatox.v  1  rrspi(ti\  rly.  Tlir  ilrtoxii-a- 
liiiii  pi'iii'i'ss  iii'i'iiis  ill  till'  Umt,  as  lias  Imtii  sln'Wii  hy  rxprriniriits  in 
whii'li  this  ()i-u;aii  was  artilii'i;ill\  piiliisi'il  milsiili'  thr  Imily.  Tlii'y  afc 
tlii'ii  n'iniivi'il  riiiiii  thr  liliiml  liy  thr  Uiiliii'xs  ami  I'xrri'tcil  in  hi'  iiiinr. 
Till'  piiipiiitiiiii  lit'  I'tlii'iTal  siilpliati's  in  this  lliiiil  is  thfril'mi'  an  imlii-ii- 
tii.ii  III'  till'  I'Xti'iit  lit'  iiiti'stiiial  puti'i'l'artiiin  nf  proli'in  i  sit  pajrr  li:'.".'). 
Till'  inilii-an,  liiiiit;  rrailily  ilrti'i-talili"  liy  thr  wcU-kmiwii  nlor  roai'tion 
nf  .lallV',  s.TM's  as  an  imliratni'  nl'  thr  extrnt  nl'  iiitistinal  putii'rai'tinn. 
Till-  iniliili'  anil  skati'li-  "liii-h  aii'  mit  thns  alisniln'il  ami  ilctoxiriiti'il  arc 
I  xi-ii'ti'il   with  till'  t'l'i'i's.  til  wliii-h  tlii'v   -riM'  tl Iiaiartrristii-  mliii' 

Till'  si.uiiT  of  the  plu'iiiil  is  tynisiiii'  1  that  nl  tin-  imlnh'  is  tiypt"- 

phalli'.  Thf  rlii'liiii'iil  pinri'ssrs  invnivi'il  an-  slmwii  in  thr  I'nllnuiii'j 
■  i|mitiniis.  in  whirli  tlw  liy-pnulmts  nl'  tin'  rcac-tiniis  art'  in  ln-arki'ts. 


coll 

//\ 

III        I'll 

roll 
111-        111 

roll 

/\ 
III-        III 

con 

11'            III 

coll 

//\ 

lie         III 

lir        I'll 

III'        III 
1  • 

III-        III 

1 ' 

lir        CM 

lie         III 

\/ 
III 

III 
CIIMI 

<-|| 
Ml,i 

(11 

III 

III 
'  II  (1) 

riMiii 

111 

C|l      .1) 

no, 

coon 


il""'M 


c(M)n 

\\  pln'I'V  I 


iiinii'  iK-i 


,   |...\V| 


VI'llt'MVl 


i  I  iii;wii'-'iil  I 


l.h.'liol, 


'} 


I'tic-  :ii'i.|  I 


.via 


Illlil.o'l'lUS 


I'lllri't'lli'l  lull     III'    tl  \  |i|i>|>lliillr     Is    |i|'iiliillil\      |iI'i'i'imIi'i|     Ii\     tli'Illlliili/lltiiili 
<M  I'll 

lir        <■        (•    I  II  I  llMi.riMill  lie        !■        !•    I  iM'ii  riHiil 


II"'        I'        <|| 

\  /  \  / 
III       Ml 

t\\\  |i|i>|iliiiiii'  I 
(11 

//\ 
III'       •'       I'    III  I  mill 


I  Ml 


III'         ••         III 

\/  \/ 
III        Ml 

(  illiliilf  llri'tii'  tii'iil  1 


III        II II. 


Ml'         I'         III 

\  /    \  / 
I'll      Ml 

I  ilhliiji'   |.|ii|>iii|iir  III  ill  I 
•11  III 

lir        I'       III  III'       I' 


III'       r       III  III'       !• 

\y\/  \    \ 

I'll      Ml-    ■  I'll  .         I'll      Ml 

in.liiln  (■•"klllnll'l 


III        II  II 


I'      I'll 
I' 


If.    IliiWi'M'l-,    Ihr   i-ailm\yl;isi>   luirlrl  i;i    HliniM'    llir   rnilmxvl    !.'l'mi|i   In 

rmi'  till'  iiiniiiii  ifriiii|t  1ms  liri'ii  iimiom'iI,  liiiriil^  tnxii-  siilistniii'i's  i-iilii'l 
jiiniiirs  jii'i'  )iriiiluiTi|.  Tlii'V  .iii'  tlir  sii-falli'ij  iiliDiinim  •<.  Fnim  aliniiiii. 
I'fliylaniiiu'  is  rniiiicij ;  I'khii  txiusiiu',  pliiiiiililh.x  laiiiiiii';  fiiuii  hisliiiim. 
wliii'li    if    will    111'    ri'iiii'iiilii'i'i'il    is   Mil    iiiipiiitaiit    |>i'iili'iii    Iniililiitifstnnr 

iiiiiila/vIi'tliN  laiiiiiii',  ami  sn  mi.     Tln'  pr ss  nj'  I'm  iiiatiun  is  illustialnl 

ill    till'   ai'i'iiiMpaii.v  iiiLT   fni'miilif: 


IL 


'i?; 


I.  CM  rill  Ml  I  rituii      I'll    ,  III  III  I  Ml  1 

AI:Miiiir  litlivliiMiiiM' 

-'   fjl,  Oil  >. I'll  rilMi     iiioii      11)       r.  ll.MilJ ,  I'll  III  Ml 

Tviiisiiir  l'l!Mi\litlivl:iiMilir 

:.  I' N  II  I'll  .ril.MI    .rituli      in       f  ii  N  .  m  i'II.mi 

Hi«'i'lill<'-  Illli.lil/vlitllviiiniiMr, 

Similar   siilistain-i's   an-    mtv   i mmi    in    tlu'    mrlaliiilii'    piiHliii'ts   ni 

lilarits:  for  cxaiiiplf.  tlirv  I'lmstitiitr  llir  artivr  priiii-ipU'  nl'  in.'nt.  TIhv 
arc  also  no  ilmilit  picrjiii-i-il  in  tjic  tissiirs  <,(  mamiimls,  iniiilazv  Ictliv  la 
mine,  oommciiily  ralli'il  liistaiiiiiir,  lu'iiiir  tlnis  iiruiliu'i'il,  as  wfll  as  tin 
.'losi'ly  ri'latnl  I'piiii'pliiiiu'.  wliii-li  is  tlir  ai-tivr  i»iiiiciplc  ot"  tlie  siipia 
ii'iial  (.'lariij  isin'  pai;i'  7:>7i.  ami  ina.v  lit'  ili'sriilii'il  as  a  iiiftliyjati'il  i'tli\l:i 
miiio  ilcrivative  of  tyrosiin'. 

I'lionylni'rti''  ariil  ))riMluiM'il  I.y  a  similar  pniri'ss  fioin  tvrosiiip  iiia> 
lie  i-xcri'ti'il  ill  111.'  iii'irii'.  wlnii'  it  I'linns  tlw  miitlnT  siilistaiu'o  of  liniiin 
iri'iitisii'  a.'iil,  to  wiiicji  tlic  ilaik  l.iouii  color  d'  i|i(.  >iriiii>  in  alkaiitoimii.! 
is  (Inc. 

Tlip  (.'I'rat  importaiii'r  atlai'lii'ij  to  iIhm'  ili'i-oinposiiioii  |iriiilni'ts  oi 
proteins  ilopfinls  on  ihc  fad  that  tiny  liavc  powrrfiil  pliarmacolou'ir 
actions.  Tlii'sc  artions  art'  ilrvi  loped  very  lari;el\-  iijion  ||i(>  vas'-M'-'o 
system;  liistamine.  for  example,  produces  marked  vasodilatation  ami 
lowers  tlio  coaiinlaliility   of  tlie  M 1.   wliereas  oilier  siibstanees  of  tin 


IIAlTKKIAI.   KKilHTlOV    IN    TIU.   INTKSTINK 


rm 


.Miiir  I'Inss,  liki'  rpiiM-pliiiiif,  liiivc  flio  prcipcify  (if  raisiim  tlio  Mooil  prrs- 
^iinv  !••  ItirKcr  lUwos,  serious  iiprvoiis  \vnipfoiiis  hikI  a  ('(uulitictii  of  pro 
roiiiul  oollnpsc  nil'  pri>(liM'<'<l.  Thoso  olwiTvatioim  havr  lr<l  scvrrni  iiivos- 
ti'.'iit<ir«  to  lu'liovo  tlint  tlic  ptTsistciit  oci-uriciicp  of  luiHcrinl  fcrmrn- 
i.itioii  ami  tlu'  nlisorpfioii  of  the  n-stiltiin.'  clecomposition  piodiicfs  of 
|.riit<'in  into  tlio  IiI<hmI  ultimately  eaiise  arferioselerosis  and  the  other  Mnip- 
t.iiiis  tliat  aeeompany  stMieseenre.  It  is  diflleiilt  at  the  prenent  time  to 
i>iiovv  how  niueh  of  this  one  w\\i\i\  to  helievi,  nlthoutfh  it  ean  not  lie 
i|niilitP«l  that  piitrrfaetion  has  nv  infavfirahle  net  ion  on  the  arteries, 
iMii   that   an  exeessive  <le|frer  i       ,  ises   the  syniptouis   of  ptomaine 

pnisonini.'. 

If  the  ptomaines  have  fori.     1  1,    '      :«■>■'    >  T         t  is  oaten,  the  symp- 
toms develop  in  from  one  t'     ■ 
turn  oceiirs  in  the  intestii'  • 


'iriie  IIS  the  food,  the  ]>'  ■ 
..luse  symjitoms  until  f  '•  !vv<i 
i\cr,  they  develop  in  ■  ■  '  nr 
iiMially  diarrhea,  whieh  ('n-'Ioj 
•  ••TeiidinR  liaeteria  and  jitoimiiiK 
Ai-fiiiil  infection  of  fond  iri'l 
iiipr  i.t  much  viorf  common  than  /. 


I  .  ii's  ;if;  r  il  i'e;i  int  if  the  docomposi- 
i>  ii^t  ■  t'  >■! '-ia  fill  re  taken  at  the  same 
.  1.  ;!■.    n>     nil -f  d         iped  suftleiently  to 

.1  ■'  r!y  i  'It  i.  irs;  sometimes,  how- 
I'ror.iii.i !  I  1'  iiontj  the  symptoms  is 

I,     i.e   ;ii;i)iosr  of  Kettinir  rid  of  the 


■■,/   III    ill     pnrnlnplioid-rntrritidis 
,1  I",      niancat  (ptonMtnrs)  thnt 


Inirr  hrrn  firnrratrd  in  food  hfforr  it  is  Inkrn  (Jordnn").  Meat,  milk 
iiiid  other  protein  ft.ods  are  usually  the  enrriers  of  the  liiieilli.  and  in  most 
r  the  aeeurately  reeoided  eases  the  meat  or  milk  was  found  to  he 
. I.  rived  from  animals  sufTerini;  from  enteritis  or  some  other  infeetion 
><.iiiietimes,  however,  perfeetly  «ood  food  may  become  infeeteil  li\ 
li.iiidliiiir.  Althouph  the  symptoms  ate  usually  aetite,  they  may  elosely 
viiiHilate  those  of  typhoid  fever,  and  the  effects  of  the  attack  may  liuRer 
I'lir  weeks  or  months. 

The  commonest  type  of  pttisoniiifj  by  sidistances  actually  present  in  the 
mod  is  that  known  as  })olulism.  In  this  the  Kastrointestinnl  symptoms 
ire  not  pronounced,— indeed,  paralysis  of  the  intestiii!)'  "ract  with  con- 
stipation is  the  rule,— hut  those  afTeetinK  the  nervous  :  feni,  dizziness, 
■lijilopia  and  other  visual  disturbances,  with  diftictdty  ni  swallowing, 
>i-e  very  prominent.  The  temperature  and  pulse  are  usvially  normal. 
Ill  practically  all  of  the  reported  cases  of  botulism,  the  so\irco  of  infection 
I  ;is  been  food  which  after  havinft  been  subjected  to  .some  preliminary  treat- 
•iiont.  such  as  smoking,  picklinpr,  or  canninpr.  had  been  allowed  to  stand 
l"r  some  time  and  then  eaten  without  cookinp.     The  Hacillus  botulinus, 

hich  is  resjionsible  for  the  production  of  the  poisons  or  toxins,  is  a 


Mr  i 


■■}       i 


■•"^ 


:.(i4 


lUtiKSTroN" 


!i  V  :f- 


n  'f  fi ;  -  i 


Ntn.M    iih.-i.iclic    ,111.1    is    ifiidily   .IcsIihn  rd    \>\    i-imkiii;;.   iis   aiv   also    li 
|HiiM.ii>       AiititHNJiis  aif    r.>i'iiir<l    l.y   siiMctlial    iiijc-tiniis.      Aliotlicr    In,- 
'"'"      '"''>      I'll''    <\ailllilr     (if    IHiisKiiiiijr    liy     |>lii(|l|cts     fnriiud     in     t'd.nl       , 
lli;il    i-aiiMij   In    ■■(•I  i^citiixiii.  " 

Tlic  licatiiiciit    ill  Mii-h  ra>cs  is  to  cin-diiiaj.'c  iliari-lii'a    liy   iri\  iiit:  im- 
irativfs.     If  the  iiiti>\i<-atii>n  is  „(  a  iiiun-  clinniif  i-liarartcr.  flic  syinpiiMu- 
air  \;,M\H',  iMiiisisi  iny  „f  ,|n.\\  siiicss,  lassitiulc.  licailai'lic,  and  jrciHTal  .1. 

1"''^^ 'I'll''    ti.aliiKiil    licic   als.i    is   to   clear   out    tlic    intestines   liv 

•jood  piii-e      Tliei-c  eaii  lie  little  iloiilit  that  many  of  the  nnliealtliy  condi 

' ^  "I'  'li''  >l<iii    leadiii'_'  to  the   t'oniial ion  of  i)iiii|)les.  aenes,  and  lioii^, 

.III'  also  laiise.j  liy  I'lii'oiiie  i  II  t  ox  ieat  ion  uitli  jirotein  <leeoin|)<isition  pn.,| 
n<'ts.      Auaiii,  |Miiyation   is  the  |ii-o|ier  t  real  iiieiit. 

''    1^  '"" "■•'Nary   in  a   work  of  this  eharac-ter  to  u'o  furtiier  into  the-. 

Iiit;li!\  iiii|i.utaiit  .|Uestioiis.  It  is  prolialile.  however,  that  the  iinportaii.-. 
Ill  the  r.'lationship  of  .•\,-..ssiv.>  protein  piitrefai'tioii  in  the  intesfin.>  i. 
iiiaii.\  of  th.'  S.I  .-all..,!  minor  dis.-as.'s  can  ii.it  he  overemphasized.  On  tlir 
.ith.-r    hail. I.    «.•    must    l..^   .ar.'fnl    ii.it    t.i    attril.iite   every   sort    of   .'liroiiir 

I"i"i'    '"    'I'i'^    pulrefa. -thill.      ToMiiiia    is   often    a    shil.liolctli    of   tl, 

prol.'ssi.iii.     Wh.'ii  a  .lir.iiii.-  .lis.Mse  .-an  not   |„.  diaL'tmsed.   it    is  pnt  dou  i 

as  ;i    toM'iiiM,      This.   Ii.iue\er.   is  Hot    1 |ie;i|   seien.-e       it    is   niiMlieal   silirk 

I"-        •'    1^   .■eilaiiil\    unsafe   at    the    present    time    to   eonelnde    that    iIm 
iii.|iiiar.\  s\  inptonis  of  sen.'se.n.-.'.  sneli  as  lirrd  .•irt..ri,.s  or  increased  lilo,,,! 
pr.ssiiie,   aiv    inxarial.ly    to   lie   atlriliiited    to   this   cause.      It    will    he   i. 
iiniiiliei.-.i   that    .Mit.-hnikoir  is  laru'clv    respoiisihle   t"or  siieh   a    view,  an.i 

als.,  that   III'  simL'este.l.  .is  the  surest   way   to  uanl   otT  the  <dia •  of  sii.! 

"''"^''••i'i"ii-  'li''  lakiipj-  of  hntt.'riiiilk.  whi-h  wonhl  supply  liacten,. 
tlir..iiL'li    nh.ise    '.'rowtli    in    the    intestine    the    I>r.itein-.|e.st  r.iy  in;:    haetei  i,: 

u.iiil.l   II. . I   lie  alile  t.i  thrive.     It   is  pmhahle  that   the  same  I'esnll  I.j  I,. 

.iilain.'.l  111  pati.nis  sli.iuni'.'  iin.loiilited  si^rns  of  siifiVriiii;  from  int.'stin.ii 
piiIr.'i'a.ti.Mi   \,\    a    i-liaii;;.'    in  .liet    in    th.-  direction   of  ^.'ivinj.'  more  earl..> 

li>. Irate,  fill,  as  we  have  seen,  if  there  is  a  plentiful  supply  of  this  f I 

siiitV  in  th.  siinill  iiil.sJin.'.  the  lia.-teria  <l.i  not  tend  to  attack  the  proteii 

I'.iloi.'  I.aviii._'  this  snl..ie.-i    it    is  interestins;  to  consider  for  a   m.nn.i 

""      '"""     "'     ""      «'■"■'      y'liiil>liii,is     lliiil     fiilloir     iiilisliiiiil     ohslnirll,,. 

Tills    .|n.sii,,ii    li^is    r.i-.nll\     I n    .liliircnfly    invest is:at.'.l    liy    Wliippl.-. 

uh..  I.. III,. I  tli.it  the  lion pr.it. 'Ill  nitroii.'ii  of  IiI.mi.I  i  pau'e  (KMii  lie.-. s  ^'r.siP 

iM.'i'.-as.'.i  in  intestinal  ol.st  i  ii.-t i.iii.  The  i-ause  t'or  this  increase  in  n... 
pi.. t.  Ill  iiitr.iii.n  is  f..iin.|  t,,  |,e  .'ni  excessive  lireak.Iow  n  of  tissue  jirotei 
.Mils,'. I  1,\  il,,'  alis.,ipti,.n  into  tiie  lilood  .,f  a  pr.iteose.  When  this  i>i. 
i,',,s.'    ix,,|,ii.,|    i'i'.,ni    ol.strn.-t.'.l    l.iops   ..f    int.stine    was    in.j.'.-(ed    int.i    fa-' 

'"-    'l"L'-,    pl..|oilli.|    N\iiipt s   of  d.'|i|-essi.in    Wi'l'.'    pl'...|llced.    tollowe.i. 

''.is.s  in   win.'li   ill.-  .|os.-   \-,:is  siilil.-thal.  hy   re.-.i\er\    in   from  twentv-f..: 


I.ACTKRIAI.   DIGLSTION    IN    Till;    INTKSTIXi: 


.■»().") 


til  forty-ciclit  ImiiiN,  Alontr  witli  tlicsc  syinptoiiis  tlu'  iiiti-ii);t>ii  t'liiuina- 
tidii  liy  flir  iiiiiu'  iiii-iciisod  In-  100  ))«m'  rout.  A  very  iiifcfcstiiip  i'iict  is 
that  aiiiiiiiils  can  In'  icndcrcil  iiiiiniiiic  to  this  proteose  liy  jjrojjressively 
iiicreasiiif;  )>ei'ioilii'  ailiiiinistratioii.  Wlieii  tliex'  are  thus  iniiniiiii/.eil. 
tlie  toxie  syinptoiiis  ilci  not  follow  upon  its  injection,  nor  arc  the  syiiip- 
toiiis  produced  liy  artificially  creating:  an  intestinal  ohstrur-tinn.  Con- 
versely, when  M  chronic  toxic  coiulition  is  kept  up  by  a  i)ai-tial  ohstnic- 
lion,  sucii  as  that  produced  by  iiiaUin;;  a  siastro jejunal  fistula  and  occluil- 
int:  the  duodcnuin,  the  animals  are  less  sus<'eptilile  than  normal  ones  to 
liroleose  injection. 

\Vc  liave  here  and  there  incidentally  referred  to  llii  run  lion  «/'  riirii)iis 
jxirls  of  till  iinslmliitt sliiiiil  roiilt  iit^.  hut  we  would  call  attention  once 
iii:ain  to  this  im|)i>rtant  subject,  especially  since  inan>-  points  of  uncer- 
tainty have  recently  liecii  cleared  up  hy  the  accurate  ohservations  of 
Lull!.'  and  [•'cnirer,'''  who  u.sed  the  electromctric  methoil  for  measuring.' 
the  li.\droi.'en-ion  coiiccntrjltion.  The  contents  of  the  duodenum  removed 
|py  means  of  the  Hehfuss  tulie  in  man  showed  n  reaction  \ar\  iiisr  from  ilis- 
liiiclly  acid  to  sliirhtl.\  acid,  dependiii;;  upon  the  |)roxiiiiily  of  the  tube 
to  the  p>  lorus  or  papilla,  this  position  beiiii:  dett'i'inincd  by  \-ray  e\am- 
iiiation.  'I'he  sli-rht  deu'ree  of  alkaliiiit>  is  surprisini;.  Lower  doxvn  in 
till'  duodenum  the  reaction  was  as  frcipiciitly  acid  as  alUaliiic.  the  de- 
cree of  acidity,  however,  beinu'  so  slivdit  as  to  favor  rather  than  retard 
the  diircslixe  jiowcrs  of  file  i)aiicreiif ic  juice. 

To  determine  the  reaction  lower  down,  the  oliser\  alioiis  were  made  on 
recently  slauirhtercd  animals  '  pitrs,  calves,  and  lambs',  the  small  intes- 
tine beiiii;  tied  olT  in  loops  of  the  iipiier,  midille,  and  lower  thirds.  The 
eontcnts  of  the  last  loop  were  often  alkaline,  but  mi!.'lit  be  more  acid  even 
than  those  of  the  first,  which  were  usually  faintly  of  this  leai'tioii.  Con- 
siderable variations  were,  however,  the  rule.  The  mi\cd  intestinal  con- 
tents of  a  recently  fed  doi.',  i'cnio\ed  immediately  alter  death,  irave 
I'll     li.TIt:  i.  e,,  very  fidntl.v  acid. 


i    Mi 


DKiKSTlOX  liKFKHKNCKS 

I  MonoH^i'aphs  1 

l':ivl"v.  .1.   I'.:      Til"  W.  rkihi:  ..r  111.    l>i-.-Ii\r  (ilini.i-.     Ti:iii>.  I.v    Sn     \V.    II.  Tl |. 

> l.iill.inll.  (Ililtili,  e.i.   J,    I'.ijll. 

■?*lioliii(;.   K.    11.:      Ke.Tiit    A.lMiii.M's   in    tlie    I'jiv^i. :!..:;>    ,,f    |>i:;,.nM,.ii.    W.    T.    Ke.-iie   \ 

'"  .  <'liii;i;;i>.    I!'ii7. 
I'liiiiiiiii,  \V.   H.:      The  M.ili.-iiiii  111   K;i.|..i-  et   lliMi>n..ii.  Int.  niiii.  Me. I.   \l. ;;i:i|.li-, 

I.eii.leii.  Kil.  Arm. 1.1.   IIH  I. 
'I'lirlsi.n.   A.  .1.:      Tli.-  riiiilri.l    el'   lliiiiy.'i-   in    ll.iillli   iiii.l    iii..:i>.-,    Iiiiv.    ..I'  i 'lii.-.i-.. 

I'r.sx.   11117. 
I'.'.l.l.  T.   Willful..:      Th.'   I'liiii.al     \ii;il..ili\    ..I    III.-  (in-.!  1..1111  .'^l  iliiil    Ti:mI.    M.-iii.li.s 

liT.  I  iii\.   I'res>,   l'.il."i. 


506 


DIdKSTION" 


(Oriv'iiial  l'iipt>rs) 

•■<-,niii"ii.  \V.  H.,  .11,. I  cm,  II,  M.K.rii:      Am.  .t,,iii-.  I'livsii.l..  IHir.,  xli.  :i!l. 
'•'•■I'si'll,  |{,:      Pi,.,..   Alii,   l'ln-<iol.  So,-..  Ami.  .Imir.   I'liVNi,.!.,   l!i|s    xh     ."i.-i't 

"'''•,.,"",."■■  "'"'    ''■    '"•    '•"''"■■'"■      ^'""■-    '"li.VH.  S...;.  .I.iiir.    l'l..vM,.r.,    l!li-,    xli^.   |.| 

-■■•Hi.l.kin,  It.  IV.  K„l,;,s,|,kin,  W.  .1..  i.ii.l  Sni.h  it.«,-l,.  \V.  W. :      Ai.li.   f    niikr.   Aiiat..MM. 

1!«'!l,  Ixxiv,  iN. 
'\fii,:illiiiii.  A.  11.:      KrpI,.  ,l,r  IMivsi,.!..  xi,  .V.is  (i.-,7. 
^'.Miller,  K.  |{.:     (/ii;iri.  .Iniir.   Kxpcr.  I'li.V!<i,il.,   I!t|:i,  \  i,  r,'. 
I'Kiikiiis,  .1.  S.:     .Iiiiir.  I'li>>i(il.,  UmmI.  xxxIv.  1:);i  U4. 
'"'K.'ftoii.    If.    \V.,    aii.l    KihIi.    r.    C:      Am.    Jmir.    I'livsi,.!.,    I!»l.-,,    xxxvii,    4<J1:    mK 

INipiclski.  !y.:     Ari'h.  f.  ,1.  j;,m.  I'hvxicil..   limi,  jxxwi,  L'l.'i. 
ii.Mclt/cr.  S.  .1  :      Am.  .Imir.    |'|i.\si-,l..    iv.iii,   ii.   ■2,\i\. 
i-CaiiiKiii,  \V.  !«.:     Am.  .((.iir.  I'livsinl.,  ISJtS.  i,  ;'„-|<i. 

1  'aiinnii.  \V.  H.,  an, I  Hlakr,  .1.  H.:      Am.  Sure.  l!"!"'.  xli.  HSH.     <  I'.   N„.  :! 
"Maialliini.  A.  H.:     S,,.  i'il/u,  nil,!.  \l.  I'.,  I'm,'.  I!„v.  S.„'.,  Ixxxiii,  M,  .".<!. 
'   llarv,v.  H    r,  II..  an. I   H,nslcy.   i;.   i;.:      Hi,il.   Hull.'.  W.mi.I 's  Ili.lc.  l!t|L'.   xxiii.  JlV. 
'■l.oni;.  •'.   11..  <■!   al.:     .lour.   Am.  <'li,iii.  S,,,-..   1!il7,   xxxix,   irc  an. I    Uir;:   als',,   il.i.l 

l!»ir..  xxx\  ill.  ,"!«. 
iMnrclnn.   K.    W:      Kii,"l    I'lii-imiim,   I'niv.  (if  riiii-.iUM   Pi,.ss.   1917. 
"•\Vhi[i|ili..  (i.   II.,  C.ok,..  .1.   v..  an. I  Si.-,irns.  T.:     .linn.   Kxp.r.  .Mi, I..   I!il7,   \\\.    i: 

.Msd  \Vlii|,|,l,-.  (i.  II..  sionr  ;iii.l  Itiriiliciiii:     II. i, I.,  liii:!,  xvii.  :.'sii  ami  :'.ii7. 
'■'l.onL'.  •'.   II.,  an, I   I'l  ni;.T,  I'.:     .Idiir.  Am.  Cliini.  Sn,..  1!il7.  xxxix.   IL'7><. 
■'■\Vlii|.|.li'.  C.  II..  jiii.l   l|.,.,|„i,  IV  \V  •      .\m.  .Ii.iir.   IMivsidl..   II'll!.  xl,  :;:;j  an. I  .';t!i;  iLl! 

I!tl7.  xlii.  LV-)7  anil  l.'iil:   11..,. |.,:      |l,i,I.,  |,.  -^so. 
-i\I.lt/,  r,  S.  .1,:     Am.  .Ii.iii.  Mv«l.  .s,  ..  l|i|7,  I'liii,  4tl!l. 


b  4  ■ 


i      I 


!      'I 


PART  VI 
rHK  KXCKKTION  OF  TRINK 


CIIAITKK   I. VIII 

TIIIO  KXr|{KT!(»N   (•!•'  I  KINK 

Hv  K.  (I.  I'l  AKtT..  I'.A,.  M.I). 

It  uill  lie  iiihisiiMi  to  introiliir.'  [hr  sulMiM-t  \>y  .1  hrii'f  if\  ii-w  «(  tin- 
N-..Mti;il  sli-uctiiijil  I'cjiliiics  III'  the  kiilncy.  in  mi  t'iir  as  tlwy  iipplx-  tn 
111'  cxi'iclDrv  I'uiK'tioii  III"  tlif  oiszari. 


STRUCTURE  OF  THE  KIDNEY 

The  kidney  is  iiuiiiiiv  ilerived  t'ldiii  tlie  Miifaei'  (if  t'le  celuin,  and  is  a 
hK'Midernia!  struefiire.  In  this  rcspec'  it  difT<'is  I'mm  ni'dinarv  scpretiti!? 
■jliiiids.  whieli  arc  eiidddeiiiial  in  (irijiin.  .hist  as  it  is  nmi'e  or  less 
iniiine  in  its  d<\  eldpiiient  as  a  ixland,  it  is  also  )iiiii|ne  in  its  method 
■  I    t'lliietiordtl!.'.     The  pliysiolonie  theories  <<(  \\,,-   nieeliaiiisni   of  nrinary 

•  iTetion  are  elosely  related  to  the  hiirhi>  eliara<'leristic  strileture  of  the 
kidney.  For  this  reason  a  inief  snrvey  of  the  stnn'ture  of  the  different 
l^arts  of  the  uriniferous  liilmies  and  the  epitii''  iai  n-ils  \\  ith  whieh  these 
.11-  lined,  is  advisable. 

The  uriniferous  tidnde,  whieh  is  the  x'eretinir  unit  I'l'  the  kidney, 
■.d\es  its  origin  in  the  ea])siile  of  Itowinan.  whieh  'nay  lie  likened  to  a 
hnllou  sphere  of  very  <lelieate  epithelium,  one  side  of  whieh  is 
.ii\  ML'inated  hy  a  very  ninidi  eonvoliiled  eapillai'v  mass,  the  (.'iomoriilns. 
riie  eapsnic  o]>ens  up  hy  a  narrow  t\\iste<l  neck  into  a  tiilmle.  whieh  is 
'.illier  tortuous  in  tin-  cortex  (the  proximal  eonxolu'cd  lulinle',  liut  soon 
lakes  a  shai'p  descendinir  eonrsc  in  the  uiolulla  towards  the  pelvis  of  the 

idiicy.  and  (hmlilcs  hack  (loop  of  Ilenjc'i   in  a  strai«:ht  ■•oursc  airain  to 

iic  coitcN,  where  it  aj.'ain  makes  a  twisicd  course  ithe  distal  convoluted 
'ihulei.  and  terminates  in  a  eollectinir  tuliulc,  which,  umtinfr  with  other 
iiliidcs.  collpets  the  urine  and  condin-ts  it  to  the  pelvis  of  the  kidney. 
Ihc  capsule  is  lined   with   \ery   thin   eiiithclial   cells,  esitccially  over  the 

.ipillaries  comprising;  the  (;lomerulus.     The  proximal  and  distal  tulmles 


i 


i  'f 


1 


! 
If 


)(!> 


INI      I.M  III.IIUN     III'    I    HIM 


iMllltilill    <'|i|tlii'lllll|l     -IhiH  IIIL'    .1     |irii|llllll|||     s|li,-llilill.       'I'lll'si-    ^t^i;|tilllls    ;: 

'""^   "'■   i-'iiiiiulis.    \\lii,-h    run    l.n\.iriU    tli.^    lunini    "(   ll rll.    l„.,-ni|.i 

sliiii't  .■|^  llii'v  ,i|i|ii,.,i,|;  It  Mini  .•i|)|);il'ciill\  Nt.iiiiliiiu'  ill  cliKi'  ii 
I'  '"  'll''  i-.itii"i  inuiiiincnl  intiTHiil  >  liiini'ii  sii-i;it|.,|  Imr,!,.,- 
I       Siiiiic  liisi,,|,,;;iviv  ImIii-m'  ih.it    til,-  ■,|ii;iliiiM>  ;il   llif   l.iinlci-  ,-, 


Ifss    ll 

I  lllll>ll 
llir  i; 


ii-i  ! 


'i.tk:*.iiii    ■  \    ll 


'■''•'">  '■'!'•'•  "I"'-!'  ■"■■  'li'-.Til.,.,!  M.  i„-n  ,^  i.Min.iiilr,     ■I'll,-  i-tiiM  ,uv  si,.. 
'"    '''-      '"'        'I'll''    'l''-i  niliiiii    liinli   ..r    Ij.'iii.V    i,„i|,    ,s    Iii|,.,|    uilli    ;i    tl 

•'■'^'■""■"'     'I'illl.   lillh,     Mill,     !,•,,■.-■     I.i.l.^lh.,;     hllrli.i.         Til.'     lll.tal     ,-.,M\nll,V 

'"''"'''  '^  '""-'I   »*''li  '-'Ii-  11. it   iiiilii^,-  ili.iv,.  i.nni.l  III  till    iM.iMiinil  liil.u;. 
-N.'.'pl   tliiil   Ih..  ii,M,-r  ,„,i,|,.,    I.  II, , I   Mii.it.,!       Tl,.'  .liaiu.'ti'i-  .,r  111..   Imi: 


TIM;    I  \(  Id  TluN    (iK    I   lilM'  509 

:'  llic  rii|isul>'    v.iiics   «itli    till'   iiclivily   of   tlii'    kiilin'v,   iis    is   slimvii    in 
I  lie  riilldw  ini;  liituifs  irixfii  li\    I'.inilii-  mihI  .Miii-kcii/ii'.' 


1.1  SI  1  \(: 

Ml 

\1  V  111  KIM 

KIIIM  \ 
M  M. 

•II  IlKSIS 
MM. 

lj:;.s 

!M..l 

liiti.it 

'!.*• 

J.I.s 

O.ll 

K.i; 

7.:: 

.'tt.ti 

MfMii   .Ii:iiii.lii    ..I    i.-ii.-iii,. 

■'     ^lnHMTIlIu-. 

I,llltti-li    "1    l.i'\itii;il    i..ti\iiliil.-  i    liil.ulf 
••      ,li-l:,l 


'I'll!'  lll'lliiiix  tllliiili'  li;is  ;i  I  ciiiMil- .ilili'  hi'niil  '.ii  jijil  ii.  '('lie  rciull  ar1««'i«'s 
;iiiM'  ilii'cftl\  rr'iilii  till'  .iliilniiiih.il  .imt.i  ami  aif  \im\  slimt.  Thi'V  run 
;hiiiiiuli  tlic  iiifliilla  til  ill''  i-tiftt\.  ai'l  ,1'iiii   \\\\\i    iii'ii;lilitiiiiii:  atlfi'ii's  In 


'I  111   an-ln's   lidiii    \vliii-li    |iri"-i'iM|    iiiiniilir ,,   lliat    lailiatf   iiiln   tl irtfX 

'I  '_'i\i'  ulV  viiialli-i-  lii-aiifla'N  .'arli  mI'  wliirli   mt>    ^lairtlv   lir-i  .ik^    ap  iiilti  a 

ai.ill  fa|iillaf\  tiil't.      t lir  irlniiifnilii^.      wlia-li  la--  in  iIh'  invairniatfil  Npliaif 

'   l!ou  man '>  fa|iNiili'.     'I'lif  i-a|iillai'ii'^  fillfat  iiiln  an  liVfrTiit   \i-.sfl.  wliicli 

|'li-;ii'N  til  lif  siiiallfr  than  tlif  .ariVn  lit   ;iilia'\.  aial   tlii^  \asM>l   in  mifi-^rini: 

'HI  till'  ra|i-iilr  aL'ain  ln-falo  ii|i  In  I'ui-iii  a  caiiillal'v  iiftuui'k  almiit  tin'  I'tm- 

liiti'il     tnlinli'-.     tnriiiin'.'    tin  i"   snii'    lilnml    Mi|i|il\.      Tlicsc    capiliarit's 

.li'.sfc  t(i  I'lirni  till'  ifiial   \rin      Tiif  liln'iii  tiT  tlu'  kiilni'>-  must,  MC'ni'il- 

-•ly.   pass  tludUuli   U\>i  si'ts  111'  rapillai'ii'v 

Till'  kiihitv   is  rii-lil>  vuppliiil  \vitli  ih-im'^.  wMfli  aif  t'ur  tin'  must  jiart 
■''!i\i'(l  iVi'Mi  tilt'  I'l'ii.-i.'  LT.'iniilinn  am!  arc  in  ■■(■nniTtinn  w'nU  lln'  siilam-li 


r.io 


Tiik  r.X(  HirroN-  np  i  iiivi: 


nil-  and  the  vairiis.  (Mlicr  lniiiu-lirs  from  plexuses  in  tlio  region  ol'  il  ■ 
supi'inTniil  linily  iind  the  aorta  join  witli  lliosr  i-oniin|{  from  the  cclii ' 
^MiiKlion  to  form  what  is  known  as  the  renal  ph'xus,  \vhi<'h  is  arrnnj."! 
in  a  network  alonj;  tlie  lilood  vessels  and  on  tlie  walls  of  the  i)elvis  ■  > 
the  kidney.  These  tiheiN  are  distrihiited  to  the  very  smallest  IiIimmI  \.s 
sels.  and  nerve  fihers  have  heen  oliserved  amonjr  tli lis  of  tin'  tiilmles 


I? 


i   1 


THE  MECHANISM  OF  THE  EXCRETION  OF  THE  URINE 

The  tireat  niimher  as  well  as  the  variety  of  suhstanees  which  are  pii-. 
ent    in   hoth   the  Mood   and   the  nrine   makes    it    appear    improliahle   llii' 
urine   exeretion    is  de|)endeiit    ui»on    ehemieal    eomhinations   within    !!■ 
renal  <-ells.  and  leads  ns  to  s<'ek  a   physiciN-heuiieal  ineelianisin   to  expl.i' 
the  phenomenon.     Can  we  diseover  the  processes  hy  which  the  kidrir^ 
faliricates  a   liiirhly   concentrated  solution   of  salts  from   a   very   dilm 
solution  of  the  same  salts  in  the  hlood  plasma?    The  ])rol)lem  is  comp' 
cated  hy  the  fact  that  the  ratios  existinti  between  the  eoneentration  ■ 
••ach  urinary  salt   in  the  urine  and  the  concentration  of  the  same  s:i'' 
in  the   lilood   are  different.      In   othei-  words,   the   mine    is   not    mere' 
■oneentrated  Mood  plasma   freed  from  protein. 
The  passajre  of  water  and  salts  thronsih  the  cal»illary  wall  and  throiiL:! 
•  liasement  mcmlirane  suiroundin*;  the  renal  cell  prohalily  takes  i)l:i' 
y  simple  difTusion.      If   it    were   otherwise,   an   expenditiire   of  enerj 
oidil   he   re(|uired,  and   it    is  diflicidt   to   understand   how   a   haseiii.  ' 
Hiemlirane  could  hriiid  alioiit  eneriry  ehanires.     Any  sulistauee  to  ;\lii 
cell    ticmlirane  is  ])ermeal)li>  will  diffuse  into  the  cell  until  an  eipi 
orinm     -    estahlished    helween    its   concentration    within    the    cell    ;ii' 
t-iat  of  !       lymiih  or  lilood  jilasma.     A  nondiffusilile  sulistaiice  will  t' 
«•-!».--     he    -ell  lieeause  it  can  not   pass  thron$;h  the  cell   niendirane.  a 
il    7  e\erts  an  osmotic  ])ressure,  it   will  also  te)id  to  keep  the  watei 
wliii       it   is  dissolved   from  ('ntcrinjr.     If  water  does  pass  into  the  c 
under  these  conditions,  it  is  due  to  the  expenditure  of  enerjiy  oppii>. 
to  and  jrreater  than  that  which  is  olTcred  liy  the  osmotic  ])ressure  of  i' 
nondiffusilile  suhstances,     I'ossilile  sources  for  sm-h  eneift.v  are  the  ]« 
sure  of  the  lilood  in  the  renal  ca|)illarics,  which  would  exert  a  force  ■ 
posite  to  that  of  its  osmotic  pressure,  and  the  jiresence  w  ithin  the  cell 
a  concentralion  of  salts  (.'reater  than  is  preseni  in  the  lilood,  and  aMi' 
exercise  a  suflicicnt  osmotic  force  to  draw  lluid  into  the  cell  aifainst  : 
(isnnitic   force  <if  the   nondiffusilile  salts.      The   jiassaire   of  the   uriii. 
c<institnents  throu<;h  the  cell  miirht  also  lie  due  to  simjile  diffusion.  ' 
suhstances  passing'  throutrh  the  cell  to  lie  eNtrndcd  on  the  other  si.]' 


I        I 


Tin:  i:x(  Ki  riiiN  ok  i  uim: 


511 


ill*'  sanio  niiKTiilnitixn  as  in  tlic  1)1(m((1.  In  tliis  case,  tlic  renal  cells 
would  net  nieii'i.v  us  a  tiller,  the  iiiiiic  having  the  same  ('(iiicentiation 
iif  each  urinary  salt  as  is  i)rpst-nt  in  the  Mood. 

A  OdMiparison  of  the  eoneentrations  of  the  urinary  salts  in  the  urine 
Mini  the  l)loo(l  shows,  however,  that  the  urine  is  not  merely  a  deprotein- 
i/ed  liluod  itlusMui,  so  that  oflier  fat'lors  must  lie  souffht  to  explain  the 
excretion.  Since  the  concentration  of  the  urine  re(|uiies  the  expenditure 
"i  much  nuire  enerfjy  than  is  |)ro\  ided  liy  the  known  pliysicai  factors, 
it  is  tfeneraJly  accepted  that  tiie  reiuii  eel!  in  some  manner  supi)lies  this 
iiiergy  by  its  metabolic  activity.  It  is  impossiiile  at  juesent  even  to 
vuimise  the  nature  of  the  process.    Two  jxissibilities  may  be  considered. 

•  •nc  is  that  the  urine  is  a  liltralc  of  the  liiood  uhicli  has  i)assed  throutrli 
a  portion  of  the  renal  epithdiinn  into  the  tidiuh's  as  a  very  dilute  fluid, 
icsemblinn  the  blood  ]>lasnui  minus  its  <'oih)idai  substances,  and  that 
liiis  dilute  fluid  is  concentrated  by  the  reabsorption  of  fluid  and  of  salts 
I'V  other  cells  of  the  kidiu-y,  and  again  replaced  in  the  blood  stieam.  The 
uther  i.s  that  the  salts  and  fluid  are  each  actively  and  iiulividually  ex- 

•  reted  by  the  kidiu-y.  Whichever  condition  is  the  true  one,  the  fact 
Hiiiains  that  the  chaiisje  in  the  coneentiation  entails  the  exjienditure 
"T  a  jrreat  amount  of  ciier;:y  <in  the  jiart  of  the  renal  cells. 

The  enertry  which  the  kidney  must  use  in  the  actual  work  of  concen- 
iiatiufr  the  uriiu"   fioin  the   lluid   of  the  blood  i)lasma  can  not  be  eiuu 

I'lited  from  a  eompai'ison  of  tlie  en titration  of  the  urinary  salts  as  a 

'liiile  in  both  the  blood  and  the  uiine.  Kach  constituent  must  be  eon- 
Hilcred  apai't.  We  can  not,  for  example,  deteiiaine  the  molecular  eon- 
'iiiti'ation  of  the  blood  plasma  and  the  urine  (by  measurin<;  A^  (l)asje 
I"      and    estimate    the    work    which    is    evpcnileil    in    pruducin'^'   the    con- 

I  ritratiou  from  the  observed  ditferenee.     On  the  basis  nf  sui-h  comparisons. 
i  "wever.  it   is  said  timf   the  excn'finn  of   KM)  c..-.  <if  nrine  rei(uires  at   the 

ilniiiiUMj   .'idO   kilofiraiimieters   ct'   work    M'ushny   i.      Mvcn    this  innserva- 

'i\e  estimate  may  be  wioiitr,  f"f  it  does  not  take  iiit( nsidei'ation   the 

l"ssibility  1h;it  the  excretion  <i|'  walei'  by  the  kiilney  rcipiires  enerir.\' 
'  \pi'ndiluie  on   the  ])art   of  the   ren;d  cells. 

Theories  of  Renal  Function 

I'or-  many  years  two  ii\al  li.\  potheses  have  iloniinated  the  teachintr  nf 
'!ic  mechanism  of  renal  function.  IJowiiian  and  Heidenhain  postulated 
hat  the  eoiistituiMits  of  the  nrine  aic  seerelc<l  by  the  vital  activity  of 
'!i'  epithelium  of  the  capsule  and  the  tuiiules.  The  irlomerular  capsule 
■  retes  the  water  and  the  easily  <liffnsible  salts  in  a  dilute  solution,  and 
■lie  uriniferous  tubules  add  to  Ibis  lluid  the  various  organic  and  inor- 
■-■iiiic    sidts   to    brinif    tlo'    mine    tn    thi-    iieeis.s.ii  \     coucentratinn       This 


i.  :    ■ 


.-.12 


rrii:  i  xnii  Tin\  m-  riiivi; 


r  .  = 


lllfiii'X  lins  liccii  li'iiiM'd  till'  rilnl  llnnrif.  I.MiIwi;;,  on  tlic  ntlii'i"  liilii'l 
,'|i|\  iitii-i'il  wli.'il  is  ti'i'liii'il  till'  iiIivsJimI  iIii'im'N.  wliirli  ImliK  tlwit  lli' 
Kliiliirniliis  ;iiii|  I'.'ipvillr  iict  siiii|>lv  Ms  :i  liltt'i',  wiiicli  .'illows  tin'  lliii>i 
lit'  till'  liliiiiil  |il;|siilil  til  |i;i'>s  tlll'iillllll  ill  :i  \  I'l'V  ilillltr  Millltiiill  illlij  II 
lill'i_'r  .'llllnllllts  Tills  lillid  is  ■■nlii-clltl'i'lti'il  liv  pliysiriii'lK'lllil'ill  pi'di'i'ssi'v 
iiii  its  |i,issii};<'  iiliiiiL;  till'  iiiiii;ii\    tiiluilrs  to  tlic  |ii'l\is  iil'  tlir  kiiliii'.w 

Itiitli  lit'  llii'si'  tliriiiit's  iii'c  iiiai|t'i|ii,'il('  iiiiii  I'.'iil  til  explain  tlic  plu'imni 
riia  wliirli  ii'si'iirrli  has  slmu  n  to  nri'iir  in  tin-  kiilnrv,  Imt  thi'y  Inn- 
si'ivfil    til    ili\iliip    «liat    Cilsliiiy    ti'iliis     (I     iiiikIi  III     llniiril     "I"     lll'iiiai'. 

I'MTrtinll. 

The  Modem  Theory  of  Urine  Formation.  This  thrmy  aicrpts  tli. 
^.'I'lii'ial  si-lii'iiii'  III'  liltiatiiiii  an<!  icalisinptiiin  <it'  Luilwis.'.  hut  pa\.s  ilu' 
rcspii't  til  till'  fact  that  the  kimwii  physical  t'oiccs  arc  imt  ai|ci|iiaii 
til  explain  the  reahsniptiiiii  which  ninst  ni'cin-  in  the  tuhiilcs.  It  thei'd'ni  ■ 
snppleiiients   l,iiilHii;'s  theiiry   liy   assiiiiiint;  a    vital   activity   "ii   the   p.iM 

III'  tl pithcliiiiii   lit'   the   tiiliiiles   in    lealisnrliint.'    Iliiiils   ami   salts    I'l 

the  ililiite  fill  I  ate  eiiniiin:  I'rnni  the  uliitnenilns  aliil  capsule.  A  lai'." 
aiiiiiunt  111'  |ilasiiia  lluiil  is  tilteieil  thiiiiiyli  the  walls  of  the  (iliiinenil.n 
\essels.  This  lliiiil  has  the  same  ciiiicciilratitin  nt'  the  salts  tn  which  tli' 
caiisiile  is  jici  iiiealile  as  dues  the  liluuil  iilasnia,  Imt  it  is  free  nf  the  c'! 
Initial  siilislanccs  iiuiiiially  present  in  the  ))lasiiia.  The  lilnml  Icaxinir  ll  • 
irliiiiicrnliis  is  tlieiet'inc  a  simiewhat  ciincentiateil  snliitinn  nt'  iilasiiia  cl 
liiiils.  iiiiil  must  have  retuiiieil  tn  it  the  jirnpei-  anmunl  nt'  water  nii'i 
sails  In  iiiaUe  it  an  nptiiiiillii  tlniil  t'nr  the  limly  cells.  This  is  accoinplish.'.l 
liy  active  alisni'ptiiui  t'rnni  the  elniiiernlar  liltrate.  The  salts  that  are  •  ■ 
III!  Use  tn  the  linily  a I'e  not  reahsnilieil  aiul  thcrefnre  ai)peai'  in  luL'hl' 
cnnccnliatcil  t'lnin  in  the  urine.  These  salts  are  teiiueil  nnntlin  •iliohl  sn'' 
stiiiiii ^,  anil  since  their  preseuee  in  the  |ilasina  is  unnecessary,  they  cni 
tiiiiu'  tn  he  cxcrcteil  as  Iniii,'  as  they  are  present  in  any  eniiecntratinn  in 
the  liliinil.  The  salts  that  arc  necessary  fur  the  jilasina  are  teriiii'' 
flrfi  sliiilil  siihstiiiii  I  .\-.  ami  are  reahsnrlieil  until  tlicy  are  .iiraiii  i)iescnt  i 
the  plasma  ill  iii)tiiiial  streiii.'t,h.  Fur  exaiiiile.  urea  ciiiitinucs  to  be  e\ 
ereteil  as  Iniii.'  as  any  is  present   in  the  liliiml,  wliile  u'lucnse  is  conipletri 

Tcalisiirlicil   sn   liiiiir  as   its  cni ntratinn    reinains  iimlei'  a    more   or  \i- 

liM'il    level. 

It   is  iiupnssilile  tn  -.'iM'  a  siiiiiiiiaiy  of  the  arirmneiits  wliieli  have  lici' 
ailvaiiceil  in  support  of  any  nf  the  tlieories      llowcvcr,  since  the  moilc 
tlicorv    ap|)iars  to  ntTer  a  lictter  c\|>lanatiiin  <if  the  estalilisheil   facts. 
iiia,\  lie  \\  ise  tn  rccniiiit  snme  nf  the  liesf  experimental  evidence  in  snpp'  ' 
nf  it. 

First,  we  must  im|uire  as  tn  tlic  amount  nf  deproteiiii/ed  Mood  ])hisi! 
which  the  capsule  must      lici'  ofT  from  the  Mood  in  order  to  furnish  ti 


Till       I    Si    111    llllN 


'.in 


.iMHiiiit   111'  till'  \Miiiiiis  s,ilt>  i\i-i  .l.il   i-.M-li   ilin    Mii'l   ill"'  .iltiMiiiit   "if  \\;ili'i- 
lisdi'linl  li>    till'  i'|iillii  Iniiii  ul   ilii'  iiiliiilrs  III  MIT  iiiiit    I'lir  III)'  I'liiici'iit  r.'i 
inii   ill   u  liii'li  till'  Mills  all'   riiiiinl   ill  llii'  mini'       In   nrilir  tu   |ii'iii|iii>i>  'JD 


ji.iliiH  111'  nii'ii   111 


I'JIMI 


!•  I'    III   iiniii'    111!  Ill' 


li.    Ill 


a  I 


llill!;  <l  IC!   |ii'i'  I'l'iil    III    Ml  r; 


,-l\    !■      I. 


:is|nii    lliiii!   riiii 

lllllliill     llic    I'.'lll 


•_'() 


n.(t:i 


lUIOIIi 


!l    lilir-   III    wall 


•  I    III   III!'    Ill I    I'liini 


iliiliit't'l-olls  lliliiiirs       SiiH'i'  ijiii  1,1 Ilji.v,    iJiiiinuli   tl,,'  \i\i\ 


III  '  s   |>   \  I'l  \' 


:ival,  :\i  Ii'ii'^l  niltl  lili'is  |h  i   .j; 


\  .    nhl  \     .1 


liiiiii    |:!  |ii'r  !•'  Ill  III  llir  iliiiil  i-iiM- 


iii    till'    Ml 


|i.i--s1|il;    tlii'iiir 


rh    I 


\v     L:iii|iii  I  iilii>     "  ml 


I.I     |i 


natiiiii    llmillijli    tlii'  i';i|i'<l||i'  ul    j'.i.  \  lii.ill 

Till'    I'iii'l    tli.lt    siii'li    a    laiiir   aiiinillil    nf    lliiiil    Nimili]  liavr    In    lii'    ii'aii- 

•  I  liiil    rimii    llii'    iiriiMl'i'iMiis    liil-uli's    i  .'ill    litris      is    a  |iii^silili'    a    jn  imi 

ilii'islil    III'    liir    liiriil.v.    lull    (   li^hiiv     |iiiinN    niil     llial  l|tr    iiii|nii|||     larli 

ilillll'    wiiuM    ll.lM      III    ali--ii||i    I"  I     lliilll     >    iilllil    |.r     \if\  ^iiiall       ill    liis    r\ 

M  riiiii'lit  nil  a  cat  aiiiniilitiiii;  In  liss  tliaii  null 

I  III 


!•  I',  MiT  Imur 


III'  filtral  imi  nl'  ttir  |n'iiti'iii  I'l  I'l-  M I  lliii.j  iliiiiiii;li  till'  ii'iial  i-apNiili 

lli.'il   tliiiiiiijli  ail}    nlliii    iiiiiiiliraiii'.  il.'|ii'iii|s  mi   scvnal    I'ai-tniN       i  I 
IT   iiiiisl    111'  a   ililVrii'i in   till'   ]Mi'>siiii'   liilw  |.|.|i    till'  111 1   jiihl    til 


liltiiitf.       111    III- 


aiinlatnlV       till'      |ili'-,>l|| 


iiM'il    ill    liltciiii'. 


I.v  sii|ii)liril  li>-  <:ia\it\-.  lull   in  tlii'  rasi'  nf  tin'  liltiatimi  nf  tlio 


iinni' 


ill  tl 


K'   rap'.llli' 


till'  f. 


funii'«li('il   liv  tl 


ll'       pIl'NSlll'l 


if    lilonil 


iliiiiii'i'iilar  \  I'Nsi'K 


iiai'ti'i'  nf  thi'   lilti'i-  ili'trriiiiiii'vi  \\liiil 


staiii'i".   sliall    |iasv       'j'lir    iriial    rapMllr    is   a    iiiriiilnani'    iinriiially   in 
vinlis   III   till'   plntiins   nf  till'   III I.   liut    )>i'l\iiins   til   till'  ntlicr  I'liiistiti 


lis.      rinli-r  I'crtain   i-niiilil  imis    it    insrs   tliis   i-liaiai'trr. 


'I'lic  clinr- 


■•'I'f  nf  till'  lluiil  ili'ti'iiiiiiii's  |iin\    ii'ailil.\    it   will  ti'ti'i-  iliiniiirli  tlio  nuMii- 
■.nil'.     If  till-  Ihiiil  cniitaiiis  a  siilisiain'i'  wliirli  i-aii  imt  pass  tlii(ii|i.'|i  t!,i' 

'll'l'    ami    wllirll    I'SI'lts    mi    nslllnlii'    plTssliri'    in    nppnsilinn     III    till'    lilt.'lillL' 


till'  rati'  nf  (iltralinii  as  w  rll  as  i 


11'  ainmint   lilti'ii 


I'll,    Mill    l.r    I  I'l 


If  f! 


ic  I'apsiili'  ai'ts  as  a 


tiltiT  it   s| Ill  l,|.  piissililr  In  alli'i'  tl 


liii-.'il, 
ate   nf 


'iiii'  iMTi'linii  li\    \ai-\in'^  aii\'  nf  lln'sr  fai'Ims,  ami 


r\pi'iini''n 


l.-illv    til 


Till'  fai'ti 
1 1  1 1  \    hill'. 


Ill'   \  aliril 


III  si'\  riii  I  I',  a^  ■ 


If  I 


ilV  ., 


'lal   I'l    llii'   liiaii 


II'    lilil.iil    pllssllll' 

if   I  111'   am  1,1.    t  III'   lliilll'   is 


iprii-ialilv    ill 


II     II    till 


pIl'SSllll.       Is      lll'i'll'.'ls 


.III-      ll\-      I'l 


inpi't'SNiiiir  till'   iriial   ailiiv    li 


\    nil-alls  nf  a   siti'w    i-iaiiip,   tl 


■ iMt    nf   iiriiK'   is   ili'i'i'i'asfil.      Ill    till'   ,'11  li(ii-iall\     pm  fi|si  il    Kiilnry.    llii' 

li'l  -Miilinir  frniii  lli,'  uii-trr  im-iiasi's  as  tl,,'  pii'ssinT  n\'  tin'  |)i'i'fiisiitii 


;i'!    's  ilii't'ciiscil.  ainl   iIiti 
aii>i('s  in  till'  iii'cssiiiT  in  tl 


as  till'  pii'ssiii|.   1^  ili'i'ii'asi' 


ll.     Wli.'tli 


I  .Hi'  ilirii-ll.\   ii'spmisihli'  t'nr  \;ii'intinns 

111'  rate  i>(  lliilll'  i'\riTtiiiii.  Ill'  wlii'llirr  tl  i'\    a,  I    iiiilii  irtly  li\    Mifviiii; 

I'a'i'    nf    till'    li'nnilllini      in    llu'    Kiiim'Ss.     Ii.-ls    liri'li     tin'    slllii,.,-!     nf    nillrl! 


514 


Till     I  \i  HI  TION    OK    I  IllVr 


i  f  ,=  i , 


n 


ilcliiltr.      I*l'iiliiilil\    liiilll    laitiils  nil'   iii\  nlvi'd,   as   i->  sliown   l'\    IIk'   fnl|ii\ 
iiii.'  ii)iMi\  iitinh--.     ir  llif  liliiiiil  |ins>iirf  In  itM-ic;is.c|  )»y  \  iiMicniisf  rirfi. 
ill  till'  N|i1;iiii|iiiic  !ii..i  |ini.|iiiM'(|  liy  vliiiiiiliili''!!  Ill' (In-  -.i-laiii'lmii'  ihtm-^ 
llir   lliiw    III'  IiIimmI    lliiiiimh   IIh'    kidney    is  dcrriMSccI      ml    till*  rxiTi'tinii    i.' 

iiiiiii'  r.ilU      .\|ipiiii'iill.\ ,  si'iTitiiiii  I'iiii  riiiitiniH'  •  III     iiN  liiiiir  lis  il ' 

lliids   I'l'   till'    |ililMllil    illi'   lint    lintillily    illi-l'iMMil     I'lir,   ils        ii'   •isiniitii'    |iri'vNii' 
dilr  III  llir   iliilill'll-.ilili'  I'lilinids   lisi'S,  till'  |>li'^>.il|-f  in   'In'  I'ii |iillM lies   is  'i  ■ 
luliuiT  ;ililr  111  ii|'|ii.si'  it       Till'  >;:|ni'  |iiiitit   liiis  lii'i'M   lii'iiiitit'iilly  sliiiw  ii  !■■. 
Slatiiii'.'   iiiid    his    |in|iils.    w  jui    rniiiid    tluit    lli      ■v,.ciTP.in    nf    iiriiii>    ci  ,im^ 

"  lii'ii  11 ii|iiliiii\    pii'ssiiif  ill  till.  mI .■iiilus  Ci'll  Im|...     tliiil  (Xi'i'li'd  Ii 

llic   iisiiiiitif   |iifssiii I'  till"   111 I    |if(itiiiis.   the  ciitiiMl    iircssiirt'   In  in  ■ 

I'riiin  .10  til  in  iiitii  II'.'  'riii'>  iiisii  t'liiind  tli.it  iliiilliii|i  .  '  ftii'  lilimd  wil' 
siiiiiii'  SI. lilt  inn  liy   ri'diii'ini;  tin-  iiMnntii-   |iirssiin'  iit'  tin    |i  .ilcins  in   tl 

[iliisiini,  -.VMS  ;i ni|i:iiiii'd  li\    iiii  Iiumimsi'  in  tin'  liiti'     f  i'mti'I  inn  .  rw 

tiiili  in  siii-Ii  r;i-i's  liiiiiij  in.iiiitiiillril  ,il   ii   lilmid   |iiTssii    ,■  liclou    llir  iniiin 
riitii-.ll   IHcssllic.       it'  till'  dilntinil  iif  tin-  lilnod    w  iis  I,    till'  v  'til  s.ilidi'  I'ui 

l.iiiiiiiL'  i.'i'ljiiiii  111'  uriii  iiiiiliii'.  fill  till'  ntlii'f  luiiid.  Ilii'  din   .lie  I'fTi'i-t  w.i 

'.'fiiilly   ili'i'ii'ii»i-i|.  ••ind  ;ili\    t";ill   in  tlie  li! I   |>ri'ssnri'  x\;is   I'lillmi  id   I'V 

sii|i|ni-.siiin    ill    t!;c    mini'    i  Kiium  itnii '  i ,      Tln'se    i\|irriiii' nl  ^    ividin' 
flidii'.-ltc    tli.il     till'    s.ilii  1'    |i|iiilii(Ms    its    dilliisis    liy    diilltill!;    llie    [iImsii 

liiiitiihN   iind    liiv,  I'liii'j    tlicir   iiMiiiitii'    ini'ssiiii'.    ^inei'    wlicii    tl sni.,t 

iMi'ssiiii'  111'  !tii'  lilitid  is  iiniiiit;iini'i|  liy  tin'  iiiMitinii  uf  ruliuids  in  wlii.  i 
tliis  is  iHi-Miit,  nil  dilliisis  i.i'<miis.  'j'lii'  siu'iiilii'iimc  nl"  tln^,.  fint^.  ;■ 
innili'l'tinli    x^itll    tlir    r.lisill'J-  "I'   liiVMTi'il    liimd    |  ilTs-.n  I'l-   Jlfti'l-   Ili'llliifllni'./ 

Ii.is  ,ilr.;hl,v    linn  .iIImiI.mI   to    I  |..|.jr   l:!;ii. 

This   \  ii'w    is   i-uiiliii 1   liy    till'  I'Niii'iiiiirnts   ut'   IViri'inft    ;ind   Stiiini. 

X\hii     shi.wid     tliJlt     the     ii\.\ '.■III     riinslllll|itii.!i     is     iiftcii     linf     il|l|)l»Ti;i!i' \ 
r.iisi'd   iliiiiiii_'  the  diiirrsis   pn  iliii'id   liy   th.-  iiijcctinii   uf  saliiii-.      Il'  tl 
diiiii'sis   jiinduii'd   liy   this   ini'ans  was   diii'   to   an   ai-liial    iiii'icasi'   in   )1 
Willie  111'  Ihi'  hiijiii'v,  till'  ii\\i;iii  i'iiiisiiin|iliiin  wmiM  have  Ihtii  iniTca-i  ■! 

In    the    I'riiir,   III,,   .jlnnniiiliis  and    I  In'   tlil.iilcs   are   su|iplieil    with    h' 

liy   the    iriial   artiiv,   as   is   the  e;ise   in    the   niaiiiiiial.   Init    tin-   tiilnilr^   .• 

finilsiv   eiiiiiivh  ;iie  alsn  supplii  d  \v  illi  SHI f  |he  III |  emniiiu'  I'mhk  il 

Inuer  I'Ntliliiiti.s  and  the  tllllik  thlMilU'll  a  vessel  uliieh  lias  III)  emint. 
part  ill  the  niaiiiiii.il  the  n  iial  pin't.il  vein.  The  hliiiid.  Iheiernre.  whi' 
is  siippliid  til  the  liiliiile  Is  a  niistiiie  rriiiii  the  '.'liiiiieniliis  and  the  in 
portal  s\steiii.      My   li'.'.il  in-,'  the  renal  vessels  it    is  imssililc  to  eiit   otT  i 

'■' I  ^ii|'l'l.\    "''  I'll'  •-'liililirilllls  while  lenvinir  Ihe  tlllniles  supplied   hv  ' 

ri  Mill   portal   vein       Voriii.illv    the  pi,  ssuie  in   the   renal   piirt;il   svslini 

not    slll'lieielit    111    I'lilee    III j    li-ek    tlirollirh    the    jl rillar    vessels.       1,1  ' 

'""•    "''   111''    renal    \  essels    lit    e    results    in    Jl     -iippressioii    ot"   the    nil 

ir    the    L'loiiiei  iilar    vessels    an-    pel  I'llsed     with     li'illii'er's    solution     a! 


li  II 


TIIK    i:XrHITH<\    (IF    I  HINT 


.i: 


|.i'i'<.siii'i-  i'i|iial  til  lliat  t'oiiiiil  ilk  ilic  iiiii'tii,  n  <'iiiisii|i'ralil*>  lluu  of  tliiiil 
iiuiy  lie  Mcciii'ol  from  (lit'  iirrlciN,  liiit  iin  lliii<l  is  •iliMtiiicil  wlicii  tin'  oiial 
|M>rtal  vein  is  pcifiiscil  at  a  |in'SMilr  I'ljiial  tn  tlial  imi  tnailv  pi'i-si'iil  in 
tills  M'iri.  I{ii\\  iiti'iM' ami  (iiin'^lilx  "  luml  that  |ihfiiiilMil|i|iiiiM'|ilithal>'iii 
[ililril  to  till-  pi'i  riiNioii  lliiiil  pas>i'il  tlii'i>u<.'li  till-  I'l-iial  (lortal  m-Iii.  ilid  imt 
Misi'  si'i'nti<iM,  Imt  w  Im'ii  iiica  was  ailili'il  In  tin-  iirrl'iisati',  lliiiil  'iiii 
iiiiiiiitf  llir  il.M'  was  iilitaiiifil  I'miii  t!  i'  ih-cIit.     riil'oi'tmiati'ly  (In'  pn-s- 

-HIT  <'lllpli>>  I'll  in  tlicsr  ('\pcl  iliHMils  lii;iy  liav  alliiWi'il  siiliic  (Illiil  t(l  III' 
'Mrri-ij  liackwaril  intn  tln'  uli'iinriilils.  si>  Mial  the  ifslllt.s  may  In-  tine  t<> 
I  llialinli   tliimi'.'li    lilt'  rapsllli'. 


I             llnu,  ,ni   ..I    1.1... I   ..ii.].K    ..I    M.ili.n.tn.in  .  .,r|.>|..  I.-    .ru!   •■(      ■>•'■■   .111..!    I^.!.iil,.    i:<    :i.n,.liil.i.,M 
1.1  I lU,   li.utii     fi'.'i.     I  :'.I|>|>^| 


ll  is  L'i'lii'nilly    ai'i'i'pli'il  llial  tin'  priMif  tint  tlic  i-aiiMilc  arts  as  a  (illi'r 
I'aiily  (■oiiipjitc.     liirditMiiali'lv    sin-li  i|i'cisi\i'  ispol  iiiinitiil    I'arts  can 

"!  Im'  iilVi'inl  til  pinxf  tin-  assiiniptiiiii  thai  lln-  rpit  iirliiiln  nf  tin'  tillnili's 
iliSMfhs  till'  rxi'i's  lit'  wali'i-  anil  sails  wliirli  aii-  lilti-rcil  ulV  tliimiLrli 
'   capsiili'.     It"  till'  nniili'i'ii  tlii'in-x   nl'  iiiinr  i-xi'ii'linii  is  n  nci-t,  tin'  i-i'lls 

■    til,-  tiiliuirs  niiist    nut    i>iil\    alisoili   lai'-'i'  aiiniinil.s  ul'  watfi-.   Init    \ht'\ 

'i~-l   asii  allow    tor  tlii"  I'l'i'iitianri-  int.)  tin-  lilooil,  I'itlii'f  coinpli'ti'ly  or 
ftially,  III"  certain  salts,  whili'  tlii'\    must    ii'.|i'rt  otlii'i's  I'litirdy. 
Wi'  lia\t'  I'alli'il  atti'iition  aliovi-  tn  ll:i'  I'ai-t  that  tin-  i.'loiin'1'iilai'  tiltrati'  is 
'■\  ihlVcri'iit  t'roiii  till'  Mi'ini'  that  is  tiinilly    pasM'd.     'riii-  mini'  cnntains  a 

■'■y  lii^rh  pi'i'i-rnta^ri'  nf  small  mnli'i'tili's.  ami  tin'  pi-npoi'tiini  in  wliii'h  tln-y 


MICROCOPY    RESOIUTION    TEST   CHART 

ANSI  and  ISO  lESI  CHART  No    ? 


1.0 


I.I 


;-  m 

•-     136 

U      lev 


IIIM 

IB 

12.0 
1.8 


1.25   iu 


1.6 


^     APPLIED  irvVIGE     Inc 


i   (! 


r.ii; 


■I'll I     I  \i  i;i  ■nii\    (II'    I  i;iM 


ni'c  prociil    is  iiiliirl\    iliri'ii-ciit    ri-uiii   tli:il    in   llic   lilnml   |il:i->liia  iii'  in    lli 
irldiiii'nihir  I'ltrjti'.     Tlii^  i>^  slmwii   in  llu'  I'dlliiwinir  ti;ruri"<.  wliirli  ij-ivi'  iin 
.•iMTiii-'i'  II         ;il  vmIui'  I'll'  till'  iifcM.  mil'  ;ii-iil.  i-lilniinc,  :iiiil  ulncnsi'  in   1(mi 

c-.f.  (iT  |ii'i.     in  fi'i'i'  III I  plasiini  ,-inii    Idll  c.i-,  ni'  nrinc.     In  tin    lliifil  ml 

limn    !■>    t;i\rii    tin'    clu'iiii;!'    in    rnnc'iil  rat  idii    wliidi    lias    ncciin'rij    in    tin 
l<iilni\ . 


if.' 


!!l!'.i 


Ii'li  (■.( 

.  n:(iri 

IN 

li"i I.e.  ri 

IM 

( 

II  \N(a   IN 

KKI. 

■  III.OOI 

( -lis  IMS 

s 

I  I'N 

I'lNi  i;  \il'i' 

IM  \SM  \  ("o\r 

MNS 

IN 

nil'  Kii'Ni  \ 

fi.:, 

,nr,r, 

o 

fiO 

fii.'    A.'i.i 
('111, Mine 

(IlLlL' 

.It 

.0.1 
fi 

22.7 

i.r. 

CI -.■ 

.1 

— ■ 

— 

lliM'c  till'  MiKiil  plasiiia  (liiiil  I'lintaincd  lull  iiii:!:!  per  cent  ol'  urea,  an 
ttic  iirinc  "J  per  cent       A U'diiiiilv.  t>  liters  nl'  ulnincnilar  tiltiatc  woiil 


■;  J 


I  ■< 

Hi 


H 

ii    : 


1 


1 

I! 


k\   : 


.     i  ii" 


S  ! 


;  1 


^li; 


lie   rc(piii-('(l   t.i   I'ninisli    llld  i-r.   nt'   ni-iiic,  (KKIOi.      six    liters  n'' 

I  (I.:!:! 
',;l(iiiiei  nlar  lilliaie  wnnld  ciaitain  il.li  'jraiiis  ni'  siiear.  I). 1:12  uraii's  nl' 
nric  aei'l.  and  24.11  yraiiis  nl'  (dilnrine.  Hiit  {nil  e.e.  nf  mine  ediilailis  li" 
vliiciise,  (l.d")  yranis  nt'  iil'ic  acid  and  0  (i  Liraliis  nf  elilnriiie.  Aeenrdiirj 
1(1  the  iiiddern  tlienry,  these  ti'^iires  indicate  lliat  dnrini;'  tlie  ])assaiii>  dl'  llie 
urine  tlii'diiLrli  tlie  liilmles  ."ilMIO  c.c.  df  water,  ti.li  liraiiis  dl'  siiLraf,  L't  'jraiu-- 
dl'  elildrine  and  (t.OliT  eraliis  dl'  niie  acid  wniild  liaxc  Id  lie  alisurlied  li; 
tlie  renal  epilli(  liuiii  in  tlie  )irii(liicl  iiui  d|'  liMl  e.e  d|'  \iriiie  (■(intaiiiiiii' 
the  edjieelit  rat  idli    L'i\  en    al"i\e. 

AliidiiL;'  the  iiidsl  ediiviiieiiiu  e \ perl liieiit s  that  call  he  nlTeied  in  slip 
pdit  dl'  the  ahsniplidii  ,<(  lliiid  and  salts  liy  the  Inhllles.  are  those  in 
wliiidi  the  pressure  nl'  'he  urine  in  the  tnhiiles  is  sliuhtly  increased  li.\ 
partial  eldsiire  t,\  tlie  ureter  .('nshn\  .  In  tliesi  experiments  the  iirelci 
df  diie  Uidiiev  is  partl\  clnsed  with  a  elam|i  and  the  ixeretidii  dlitained 
I'l'dm  this  kidney  is  eompared  x.itli  that  nl'  the  oppnsite  lidriual  kidne\. 
In  yciieral.  (ilistriictidii  dl'  the  ureter  results  in  a  decrease  in  the  amnunts 
(it'  water,  elildlide  and  nrea  excreted,  i'.ul,  clllidllsly.  the  lirea  cdlltellt  is 
ileereased  relatively  less  than  is  the  (dddiiile  and  water  edlllelit.  These 
results  can  he  eN|ilaiiie(l  (ill  the  liasis  that  any  pressure  aetimr  tn  (ipposc 
the  head  dt'  pressure  prnduciiiu  liltratidii  in  the  ehniicrnlils  will  rediic' 
the  aiiidinit  (if  the  '^Idiiierular  lilt  rat  inn.  and  accdrdinulv  the  time  alhi"  ed 
\'n\-  the  jiassaue  df  this  (iltrate  ahinu'  the  tillniles  is  increased  ami  ahsnip 
linn  hecdiiics  iiKire  cdinplete.     Since  urea  is  jirdliaMy  mil  alisdihed  at  all 

and    (dlldl'ide    is,    the   discrepance     In    the    eti'eets   (in    the    exeletidii    dl'    iil'c.l 
and  (di'diine  in  the  ]iai'tially  nlist  ructeil  kidney  can  he  c\]ilainc(l. 

When  \cr\    laryc  aiiidUiils  dt'  water  al'c  taken  li\    ludutli,  it   id'teii  liap 


TIM.  ixriiirius  III'  iijim: 


r)i7 


iM'iis  tliiil  till'  iiiiiii'  cxcii'lfil  ll;l^  ii  ciiin'cnt  niliiiii  til'  >;ills  less  tliiiri  lli.Ml 
|i|-cscll1  ill  llic  lllliil  111'  (lir  liliMiil.  Snliic  ill\  est  ilTJltnls  lii'lii'VC  lllilf  sUrll  ;i 
riillilitilPII     is    |)iissili|r    Lilly    mi     l|l|.    ;issUIII|ltiii||    tllllt     WiltlT    is    i|ct  i\('l  V    l'\- 

crctcd.  liiit  :i  iimmt  |>l;iiisil.|i'  i'\|)l;iiiiil  imi  li.-isci|  mi  llii'  iiiiiilcrii  llicur.v 
is  tliiit  tlic  w.itcr  tliiit  is  ;ilismlii'(|  irmii  liir  .iliiiu'iitjiry  tiMi-t  I't'.'iclii's  tin' 
l-.iiliicy  as  ;i   liiliilc  siijiiii'  sjiliiiimi,  jhhI   is   lapiijlv    liltrriil  ufV  in   ii   t'm'iii 

siiiiii'W  lial  111  I  HI'  ililiilc  tliaii  tiir  ii|iniiial  sulntimi  w  iiicli  lil I  |ilaMiia  Mil  1st 

liaM'  t'ipl'  till'  w  I'll  lii'ili;;  of  till'  tissiH's.  Tiir  tiilnili's  ri'alisniii  the  aiiHMllits 
III'  \\ali'i'  aiiil  111'  lliiisi'  salts,  siirli  as  rliluriiji's.  iiiii-  ai'iil.  aiiil  siiiiar,  iici-- 
i'ssar\'  til  iTstmi'  tin'  plasma  In  *lir  ii|itiiiial  I'lun-i'iit  rat  inn,  lull  ilii  lint 
alisiirli  till'  imiit  liii'sliuM  siilistaiiiTs,  siirli  as  urea. 

It   is  iiiipiissilili'  to  aiial.wi'  tin'   I'mi-cs  tlial   aif   ii'spniisililc    Tni'  sucli   a 

iliiii il'  all  nrptiiiii  li>    till'  ipitlii'liiiiii  nl'  tin'  tiiliu!i'~.      l-'m-  tlii'  present 

\M'  must   elassil'\    tliem.    I'm-  \\aiit   nt'  a   lietter  term,   as   vital    fm-ees.     Tin' 

.pii'stimis    that    await    inn liate    iii\  esti'j-at  imi    are    wlietlier    alisi>i'])tioii 

acliiall.x    takes   plaee.   ami.   il    it    ilm's   so.   wliat    faetnrs   eaiise    it    In   \ai'>'. 

Manx  attrmpts  liaxi'  lieen  mail',  li>  ilesi  rn\  iin.;  tlie  eap-iiles  ur  tlie 
tiiliilles  liy  means  nl'  pnismis  m-  li\  nprratimi.  tn  iletermiiie  directl.v  nr 
iliilireetl,\    the   i|llesliiin    iil'   tile    riinetimi    111'  the    tlllillli's. 

Ill    slieli    eNperimellts,    IniHever,    the    llUlillier    nl'    I'aetnis    in  \iil\"i'il    eiill 
I'lisr   the   is>iii'   anil   make   the   resiills    piaetieall.x    valueless   sn    t'ar   as  ile 

i.'i'miniiii;    the    imrmal    I'unetii f    the    tnluiles.      Other    experimenters 

have  atlempteil  tn  slmw  ahsiirptinn  in  the  tnluiles  li.\  in.ieet  iiiji-  ililVilsilili 
siilistanees,  siieli  as  ehemieals  anil  <l.\es,  intn  the  ureter  iiinier  what  the> 
'leemeil  siil'tieieiit  pressure  tn  I'mer  the  siiliitinii  intn  the  tiiluiles.  ami  h.v 
an  examiliatinll  nl'  the  lilnml  nl'  the  tissiii's  In  ileteriiiine  whether  or  nil! 
ihe  in.jeetcil  siihstanees  hail  lieeii  al'sni'lieil.  The  results  nlitailied  Iiy 
'his  mellioil  are  not  emn  ineinii,  pmliahlv  ehiell.v  lieeaiise  nl'  the  ilit'lieiilt > 
111  reaehinu'  the  tuluiles.  Ilnleeil,  it  is  \  er,\  ipiest  iiUialile  wlietlier  it  is 
pnssilile  111   in.ieet    a   slllistaliee   intn   the   tlllinles    rrniii    the   meter. 

Veai's  ayn   1  leiileiihain,  the  expmieiit    nl'  the   vital    tl iv   of  exeretimi. 

lieliexeil  Ih.i;  he  hail  ilemniistialeil  the  aliilitv  nf  the  renal  eells  to  ox 
iiete  (l.ve  siilistanees  injeeleil  int  ra  V  einillsly .  .><iliee  he  faileil  to  (iinl 
•v  iileliee  nf  live  exel'etioll  ill  the  eapsllle,  hut  follinl  masses  of  il.\  ('  ill  til" 
lilluiles  anil  staineil  urannles  in  the  eells  of  the  tnliiiles.  he  eniiclmleil 
:hal  the  eells  i>\'  the  lllluiles  Innl  the  power  In  exeiele  the  ilve.  ami  from 
:iii,iIiil;>  he  liilieveil  that  the  tlllinles  mUst  likewise  e\erete  lllC  Wilier 
ami   the    various   nrinar.v    sails,      Siiliseipieiil    work,    hnwever,    has    faileil 

In  I'ontinii   his  lielief  that    tl apsiile  is  not    I'liiieei'iieil    in   the   exeretion 

nf  the  live,  ami   il    is  as   reasnnalile  to  explain   the   resiills  of  the   expei'i 
iiieiits    with    Ihe    lives    liv     assiimili'j     Ihat    the    mass.'s    of    ilve    slllis'    iiees 
I'oiiml    ill    till'   lllluiles   ami    in    the   eelU   are    iliie   to   ihe    realisiiri)tinii    of 


:  i 


,iM^ 


')18 


Tin:  i:.\(i(i;Tii)\  of  ikinf. 


!i1m 


-:V1  -       i 


wjittT  iiiul  iii'ilijips  lit'  sdiiH-  of  till'  ilyv  fi-i)iii  |li|.  diliitt"  >.'l<iin('nil;ir  (ill  nil. ■. 
as  til  iicccj)!  Ilciiliiilijiin 's  li\  ])iillH'sis. 

Ill  till-  fiilliiw  iii^j;  talili-  taUcii  triuii  Ciisliiiy  tlif  iiKivi'iiiciifs  of  llie  cnn 
stitucnts  of  till'  i)lasnia  iiia\  In-  fulliiwul  tliiouu'li  llic  kiiliicy.  Tlu-  iilii 
male  di'stiiiation  of  each  is  iiulicatcd  in  tlie  ciiclosurcs. 


WatiT 

C.llni.ls 

I)i'\tri  SI' 

I'lir   .'li'i.l 

Si '■limn 

l'"t;is-*iiirii 

r|,I,,ii,lr 

rrcii 

Siil|ili:il,. 


I  III  MTi:i:s  rr.ASMA 

I  roNTAIX 

I 

I    I'KU 

I    CKNT  TdTAI. 

'.'■2  re'  1. 

,    s  ,-i:;r,ii    ^m. 

|~aT  i;7'  i;iii. 

j   (\.iM\-2  I.:;  •■ 

:    (I..".  :.'ii(i      '• 

i    nil;;  I".:;    •■ 

:    o.:;7  lms 

0.1 1;:  :.'ii      " 

i    (i.dii.',  i.><    " 


lij  i.iTi  i;s 

KII.TKATK 
COXTAly 

i\  Ai.r, 
lij   1. 

li;      ^'in. 
Jill  I       " 

1. ■'..:•,  •' 

LMS       " 

I'll     " 

1.S    " 


I'll 

I.ITKI 

s 

ii:Aiisiii:iii;i) 

FMIll 

(1 

STAIN 

rn! 

I'KNT 

■I 

ilTAI. 

Ill  1. 


(I.I!  ('.7     j^iii. 

ii.oiil.'!  Il.s    '■ 

0..".L'  l!l(i,.-i     " 

(MH'.l  II.S     •' 

11.111  tic'       •' 


1   i.nii:   ri;i\i: 

ClI.MAI.SS 

I'll: 

ry.sr 

TOTM 

'.<-> 

!C)0    I'.r. 

— 

— 

d.".") 

O.d.l  ylii. 

(I..!.-) 

0..1     '  * 

1 

0.1. T 

1  .->    • ' 

1 

(l.li 

(l.ll    " 

L'.n 

L'.ll     " 

n.iK 

IS      " 

n'roin    t'ii>lii;v.-i 


Tt  "i''  0  iioti'tl  that  tlic  dextrose  alone  is  eonipletely  alisorlied,  ainl 
that  til.  urea  and  tlie  siilpliate  are  not  ahsoilied  at  all  froiii  tlie  udmii 
eiiilar  filtrate.     The  other  salts  are  jiartly  alismlied. 

As  already  iiieiitioiied.  15a  •erol't  and  Straui)  lu.ve  shown  that  ili. 
diuresis  whi<'h  results  from  the  in.jeetion  of  saline  into  the  Mood  is  n..' 
aeeomi)anieil  hy  any  ineiease  in  tiie  oxygen  eoiisunii>tion  of  the  kidiie;, 
This  oliservation,  eoui)led  with  the  fact  that  tlie  total  amount  of  ehliir'ij'  . 
urea,  and  sulphate  wliieh  is  excreted  dui'in<r  saline  diuresis,  is  gi'eater  than 
under  normal  eonditini's  indicates  tl'at  the  excntion  of  these  salts  i~ 
not  due  to  any  vital  secretory  i»ower  of  the  kidney,  liut  rather  to  factm-- 
tliaf  are  extrarenal  in  oritrin. 

The  diuresis  produced  liy  addin<.'  urea  or  sodium  sulphate  to  the  liloml 
on  the  other  hand,  is  acecmpanied  hy  an  increase  in  the  oxygen  con 
siimi>tion  of  the  kidney.  This  increase  can  not  he  due  to  active  elimin.i 
tion  of  these  salts  liy  the  tuhules,  the  work  of  wliich  requires  oxyu'en 
fo?'  no  increase  in  oxygen  consumption  accompanies  the  increased  ex 
cretion  of  the  same  salts  under  saline  diuresis.  Sulphate  and  urea  an 
nonthri'sholil  snlslinices,  and  are  not  alisorhed  liv  the  tuhules.  Tin 
explanatiiui  of  tlie  oxygen  consuniptioii  is  luolialily  that  the  osm.iii. 
pressure  which  these  Indies  in  the  gl.  mcruhir  filtrate  exert  makes  i 
necessary  for  the  epitlieliiim  to  opimse  a  greater  ahsorliing  force  to  cnn 
eentrate  the  urine,  and  hence  a  gi-ater  expenditure  of  energv  is  rcipiinl 
Diuretics.  The  action  of  the  xanthine  eoniiiouinls  .'afTeine,  thc" 
bromine  and  theoiihylliiie     in  the  production  of  diuresis  is  uncxplnind 


Tin:  i;X(ri;tii)N'  of  irink 


519 


It  iiiiiy  1)0  duo  ill  i>ait  tn  vasculiir  chaiijies  and  in  i>ait  to  reduction  in 
ihf  iTsislaiico  to  liltiiilioii  liroiitrlit  alioiit  l)y  alti-ration  in  tiu-  pcrnica- 
Mlity  of  till'  capsule. 

Acconliii}.'  to  the  iiiodirii  tlieory  tlie  polyiiiia  in  diahetes  is  ]irodnced 
liy  the  excessive  aiiionnt  of  \vatei-  taken  and  liy  the  inability  of  the 
kidney  to  eoneeiitiate  thr  mine  a^'ainsl  the  osinolie  |)i-essnro  olTered  liy 
ilie  eoneentraled  suirar  solution  in  the  tiilnilcs.  The  presence  of  the  liy- 
|icr<:lyceiiiia  in  an  aiiioiinl  liiylier  than  is  pi-eseiit  in  tlie  optimal  Mood 
|ilasiiia  in  this  disease  makes  siii:ar  a  nonthreshold  siihstain'c.  so  to  sjieak, 
;md  none  is  ahsorlied.  The  'inresis  follo\\iii<x  the  injection  of  sujiiir  is 
ilieicfore  of  the  same  ty)ie  as  that  in'odnecd  hy  sulphate  and  iiiea.  The 
liinretic  action  of  the  iliu'itaiis  u'roup  is  dependent  U|)on  its  iiinuence  on 
imC  circulatory  system.  If  the  circulation  is  alieady  sullicieiit,  digitalis 
ildcs  not  cause  diufcsis.  The  ca'ise  of  the  diuiesis  pioduced  liy  iiituitar.v 
I'xtract  is  not  know  n.  It  may  lie  owiiiir  in  )iart  t<i  its  action  on  the  cir- 
lulation  and  in  ))ait  to  a  direct  action     ii  the  kidney. 

Albuminuria.  The  plasma  jiroteins  ordinarily  do  not  <ilitaiii  entrance 
into  the  tnhules  of  the  l<idne\ .  In  disease  suidi  as  acute  nei)liritis  and 
.•.irdiac  failure,  the  plasma  colloids  aie  lilteicd  olV  throuH:li  the  capsule, 
inoliahly  liecause  of  some  chaii(fc  that  has  occurred  in  the  permeahilil.v 
..f  its  memliraue  due  to  inllaiiiniation  nr  asphyxia.  In  thi'se  cases  the 
iiriue  is  usually  re(luc(il  in  anmuut.  i'rolialily  there  is  no  purely  <rl<nn- 
c  rular  or  tuhular  t.xjie  of  nephritis,  lioth  structures  sharinir  in  the  dis- 
iihility.  While  it  can  not  lie  said  that  any  of  the  so-cidled  renal  tests 
that  ha\e  lieen  advanced  in  re<'cnt  yi-ars  are  free  from  ciilicism,  tliey 
nevertheless  have  contriliuted  very  useful  information.  The  fact  tliat 
Mie  kidney  of  the  chronic  nephritic  excretes  a  urine  of  more  or  K-ss  tixed 
'o\v  siiecitic  };ra\  it.v  \\ouid  siiiry:i  st  that  here  theic  is  an  inipaiiiiieiit  of 
tlie  rcsorliinjr  mechanism,  and  the  failure  of  a  kiilney  to  exiTete  the 
[■roper  amount  of  dye.  as  in  tlie  |)lieiiolsiilphone|>htlialein  test,  sujruests 
III  im))airment  in  the  lilteriiiir  a|ipati;tiis.  Hard  and  fast  rules  can  not 
lie  apjilicd.  however,  and  prolialily  the  tests  must  at  present  lie  inter- 
preted for  the  kidney  as  a  whole. 

The  Influence  of  the  Nervous  System  on  the  Secretion  of  Urine.  In 
^liite  of  nnmcroiis  and  repeated  attempts  to  demonstrate  that  a  nervous 
iiiechanism  <;overns  the  excretion  of  urine,  no  juoofs  which  are  alio\e 
•lilicism  have  lieeii  forlliconiimr.  Stimulation  of  the  splanchnic  nerves 
'(■suits  ill  a  diminuiiou  in  the  excretion  nf  urine,  ]irolialily  liecause  of  a 
liminntion  in  the  lilomi  supply  i  f  the  ii'nal  vi'sscls  owiiiir  to  the  vasocon- 
-Irictioli.  Stimulation  of  the  \a<;us  nerves  Iniow  the  le\el  of  the  cardiac 
iraiiches  has  lieen  s.'iid  to  I'esult  in  the  ail'-rmentation  of  the  rate  of  urine 


■\eretioii 


(Asl 


ler  aiu 


IP 


L'c'-i.    The  results  are  doulitfiil,  however,  since 


k\ 


i 


WT 

r 

<!'       i 

i ' 

V-     ■ 

1 

■  -1 

! 

• 
1. 

1^1, ' 


I' ■  ' 

1 1 


'i' 


1: 


Br 


I  J 
I 

I  I' 

I  • 

I' 


<-;!  ^ 


fti 


-; 


|-q 


hfv 


■)•_'(  I 


I  in:   I  \(  i;i  I'lips   IP)-   I  i;i\i; 


tliiTc  In  iki  iiicii';iM.  ill  till'  cixyjicii  iilis<ir|iliiiii  iiiiilrr  llic  iiliovi'  ciiriilit in:  . 
'  I'l'iliv,.  ;in,|  (  ■.irl.'i'    1         h,   tlir  liuflil  ,,|-  III,.  hhhIi'I'ii  \\\v„vy  tliis  \;i-r;il  iliiii. 

sis   would   lif   ilil.'i|iirli'il   ;is  dlli'   ti>  ;|||    illliiliiti I'   till'   iilisi(|-|)t  iiill    ill    ll  . 

tliliiilcs  i-;illici-  tli.'iii  ;iii  Miimiiciitiitinii  In  tlir  :icliiiil  cxcict inn  iil'  urine. 

Tlicif  is  IKI  ilmilil  tli.it  till'  iriiiil  iici'M's  pr'nriiiiiKJIy  iil1'i'i-t  llic  ,-\crfl  i..i 
"I'  Hiiiii'.  liiH   Ih.-il   llii'v  iJM  sii  i!ir.'i'tl>    is  \v\-\    iiii|iripl);il)lr,  sii in'i-rci-i  K 


ililiM|li;itr    ivii.ii    fiilM-linii   ,-;iii    |„.   iii;iilit;iilir.|    in   ;iiiiMi;ils   liiat    li;iVi'   li;|i|    tl; 
kiijii.'ys  i-iiliici\     ifiiiu\i-,|    Mini    llini    ic|iliii-ii|.      'riici-i.   ;u-i-    iiiiiimtiuis    p. 

tii'Xcs    tll,-||    ;ilVriM    ill.'    imI,'    ,,\    mi \i-|l'tin||    liy    i sllirlidil    r.f   iji,.    rcii, 

M'ss.'In^      Injury   In  tl,,.  M.iiMi  r  nr   iiivlrr.  ,'il>i|oiiiiii;il    iiijiirii.s   li,  1  li,.   ki, 

'"■>■    '"'    '•^'•11    '-"I'l    ;i|.|)lic,|    In    III,.    -,|,ili.    l||,-|\     I, .Mill     j,|    il|,.n||||il,.t,.    MippiT 
sioli   111'   the   iiriiic. 


CllArTKI!   I. IX 


TIIK  A.Mdl  NT.  (  ((.MI'OSITIOX,  AND  (•|IAI{A(  TKIJ  0['  linXI-: 
I'.v  H.  (i.  PiAun:,  i;.A..  Ml). 

Ill  flit'  cli.llitris  oil  (liucstiiin  illlil  llictllliiilisiu,  UC  llUVC  fi)tln\\('(l  tilt' 
iiiMrsc  wliicli  riiiiil  t;ik('s  willi  csiici-iiil  rciVrcnce  tn  tlic  nutriticin  of  the 
liody.  Tlic  I'scit'tioii  ot'  tliof  cli'iiii'iits  of  iiiitiif ion  is  tiilvcn  up  uiidi'i'  ji 
iiiiiiilii'i'  of  till'  siiI>ili\i>ioiis  of  iilivsiolojix .  vi/..  r('s|iii;itiiiM,  (liu:<'stioii. 
I\ii|ll('.v'  ftllli'lioii  ;ilii|  till'  skill.  Ill  tlic  cllilptcls  on  (li>r('stioii  iittclltidll  AVils 
riiilcil  to  the  fiict  flint  flic  fccrs,  licsiilcs  coiitiiiiiiii;;  flic  iiiilittcstihlc  rcsi- 
liiic  of  flic  ,'iliiiiclit.  coiitiiiii  scvci'.'il  cM-ictoiy  |)|-(p(1ucI'  wliicli  ill  one 
!iiiic  or  iiiintlicr  have  iictiiiill>  liccii  within  the  lioily  iiropci'.  These  iii- 
iliiile  noiiii;ill\  the  piiriiiciits  of  the  liody  and  many  of  the  hcaviei-  inineial 
~a!ls,  siieh  as  iron,  niamiesiniii,  liiiic  and  phosphates;  and  under  aliliorniiil 
ronditioiis.  as  xvlicii  tin'  metals  arc  u'iveii  as  medieiiie.  hismutll  and  mer- 
iiiry.  The  respiratory  system  excretes  most  of  the  oxyjreii  and  enrlion. 
Ill  this  (diapter  \\i-  shall  take  up  the  manner  in  which  the  hody  rids  itself 
III'  the  nil  royeiKMis  and  some  of  the  mineral  waste  materials.  Kven  iit 
the  risk  of  repetition,  it  will  he  ad\anlai;coiis  to  reeapitiilalc  certain  f:i''ts 
loiK'erniny:  the  essential  chemical  stiiictiirc  of  the  urinary  constituents. 
Ml  that  we  may  lie  in  a  position  to  appii'ciate  the  kidni'y  function  in 
licaltli  .•llid  disease. 

We  iioxv  know  that  the  kidney  docs  not  form  any  of  the  s))eci(ic  con- 
■-litiieiits  of  its  secretion    i  c\ccpt    hippiiiic   acid    .      These   siilistaiiees  are 

iiii'i I  in  the  \arioiis  tissues  of  the  liod>,  and  are  liroinrhf  to  the  kidnc.\s 

l>\  the  Mood,  where  they  are  eliminated.  Hut  while  the  constitiients  are 
iiichainred  in  chemical  composition  in  the  urine  from  that  in  wliieh  they 
.111'  found  in  flic  liloo<l,  they  do  occur  in  meatiy  cliaiiy:ed  jirojiorfions. 
Ii  is  this  variation  in  the  concentration  of  the  urinary  constituents  in 
Mic  lilood  and  the  urine  which  prcscnls  the  most  important  and  at  the 
^iiiiie  time  the  most  difticiilt  (|Ucstion  in  the  plix  sioloL'y  of  the  kidnc\  , 
III  the  following  lalile  the  pcrceiitacc  composition  of  the  lilood  plasma  is 
i-Hiipared  with  that  of  an  axcraLrc  sample  of  human  urine.  The  thin! 
iiillliili  uives  llie  idiaiiiie  in  coiiceiil  rat  ion  which  ea<di  coiislitucnl  uiiilt'i'- 
■  lies  ill   p.-issini,'  I|iiou'j:Ii  the   renal   liltcr. 


Ir   !■ 


I  >-:f. 


:>-2 1 


i  I 


iSh^ 


Tin;  ixtKiTKiN  (iK  rijiNi: 


I1M"I1I  IM,\.S\I  \ 

ikim: 

CIIANGKIN 

TKIiCKM' 

I'l  l:  (KNT 

'...1 

<OMFXTIl,\Tl. 

WMrr- 

1)11  !i;! 

I'r'Mciiis.    fii 

U  nii.I 

..iImt   .•..II.. i.! 

<              7!i 



.^ 

Hivtn.si' 

(1.1 

— 

.^ 

Vun 

(i.(i:; 

n 

60 

I'll.-     !„-i.! 

(i.Mii;,' 

O.d.'l 

23 

('li;iliiiilic 

AniniiMii;i 

(I.IHll 

not 

40 

Snililllil 

<i.::j 

0,".") 

1 

l*tttM*.siiitn 

II. (ij 

(i.n.-) 

7 

(•.■.Irililll 

II.IIIN 

o.ni.T 

2 

Mii^nrMiiiiii 

ii.nii:.'.") 

ll.llllli 

f> 

ChlolilK- 

n.iMiii 

(I.J7 

no 

I'li"s|.Ii:itrs 

I'd,, 

(i.dii:! 

0  IS 

«o 

Slll|.|i:ltis    (S(),l 

Alniiin  licida 

hf 


iiflM 


The  Amount  of  Urine 

'I'lii'  iiiiMiiiht  iif  urine  pMsscl  in  t\\  ciitx  t'diir  lioiiis  vjiiics  witli  tli. 
iiniDiint  III"  (liiid  inu'i'Nti'cl  mikI  tlic  ppopoitimi  of  lliiid  rcfiiiiiiMl  liy  tin-  1hi.1.\ 
or  I'xi'i-ctcil  liv  (itlii'f  clijiniicls.  I'ikIit  oi'iliiiiirv  coiKlitiiuis  a  t«  I'lity-fcmr 
liiMir  s;iiiip)('  aiiioiints  to  from  Ktfllt  to  ISOO  c.c.  of  iirino.  On  a  constani 
"ati'f  intake  tin-  voliinie  of  urine  Is  extrcnielv  \arialile  for  any  sin-;!' 
•  lay  or  p:irf  of  tlie  day  (Addis  and  \Vatanal>(''~i.  Tlie  average  voliiino  of 
urine  exereteil  liy  twenty  indix  iiliials  on  the  tliird.  foiirtli  and  fifth  da\ - 
of  a  constant  diet  in  \\hieh  the  fliiiil  iiital;e  was  2,070  i-.c,  varied  froiM 
1.01:1  to  1,712  e.e.  for  a  twcnty-foiir-honr  period,  fr<  m  l)S4  to  l.Hl'i  e c 
for  the  first  twelve  hours  of  the  day.  and  from  fiOl  to  7SS  e.e.  for  tli. 
first  ei<j:lit  hours  of  t!;e  <li,y.  In  normal  snhjeets  the  iiinoiint  of  iiriii. 
excreted  diiriiiu'  the  iiii.'lit  '\<  usually  less  than  that  diiriiif;  the  day.  Thi- 
is  si;eh  a  constant  (iiidinu'  !hat  in  eases  where  more  than  ."lO  \wr  cent  ot 
the  urine  is  exi-reted  in  the  twelve  liours  of  the  iii^ht,  suspieions  of  rerur 
disease  should  lie  aroused. 


I*]  .  ;i 


3*.'!       ! 


The  Specific  Gravity  of  Urine 

In  urine  collected  at  ditTeient  times  of  the  day  the  specific  jrravity  iiia> 
show   a  variation  of  ten  points.     Indeed,  the  specific  };ravity  of  the  iiriii' 

has  hecn  taken  as  a   fuii<-lioiial  ti'st  hy  clinicians.     With  a  constant  f i 

and  water  intake  the  variations  fiuiiid  in  the  specific  <;ravity  of  saiiipl.- 
of  iiriiii'  taken  at   two-hour  periods  in  normal  and  pathohnrie  condition- 
are  very  useful  as  criteria  of  the  fiinetional  state  of  the  kidney.     Fix.i 
tion  of  the  sp,..-ii;,.  ir,.|i\  i)y  at  either  a  low  of  a  hi«h   lisure   is  not   th. 
usual  normal  findinir.     The  follow  im,'  fij;iiros  will  illustrate: 


AMOI   NT,    (  I)\IC1PS|||((\.    wn    (IIAKMIIU    DF    1 KINT. 


r)2:J 


DAY 


s  III        KMJ 

\  \I.  A.M. 


2  I  )  ('. 

P.M.        r.M. 


II  S 
r.M, 


I    NIOIIT 

l'.M.-.\.M. 
l.llL'd 


\nMiuil    |uT»(m  I  l.iili'i       l.ol'.l       l.iilJ      Lull       l.i'UO       l.iiUi 

III    llv|.cilrii!<i\i'    Nr|.|nlli<  |l,n|0       l.(iii:i       l.mn       limii       I, Hill       l.Hlii;    I.IIIMI 

In    Mv.H'iinliiil    I) »|.ciisMti..M    I.dlS       l.HL'il       l.iil'.i       |.ii|>       iinjil       Idi'T     l.ic'J 

ir.r'n|.il.  .1    I'll. II    Ml.   .  ntli.-il  ■»   li;.'iii'.  M.  I 

Till'  pri'lml'tioii  III'  w.'itiT  to  liitjil  'ills  is  iij'ti'ii  vi'iy  siinihir  in  iiliisiiui 
iiinl  iiriiii',  liiit  wlii'ii  Miller  is  t;il<rii  in  l;ir'_'i'  i|ii;iiititii's  llic  urine  sliow:: 
iiiiicli  iri'o.ili'i-  clijiiijii's  tliiiii  (liM's  tlic  liliinil,  iiiiil  llic  siijids  iiijiy  sink  to  a 
Miy  law  ciiiici'iitriiticiii.  (In  tlie  (illii'i-  liiiml,  u  In  ii  liltli-  tliliil  is  t;iki'ii  iir 
wlirii  tlic  skin  mill  Imwd  cjiniiiiiid'  :i  liiriri'  iiiiiipiint  nt'  llniil,  tin-  iiriiii' 
iii.iy  licniinc  very  ciiiiiTnt liitcil  willii'iit  ;i!i\'  cliiiniic  ill  llii'  Mninl  plasiiin. 
Till'  tiitiil  siilids  in  iiiiiir  ('.'in  hi'  ilctfi'itiini'il  with  .ippinsiiii.'iti'  iiccurai'v 
!.y  iiinltiiilyiii!.'  tin'  List  twn  (iiiiirrs  nf  tlir  s|ii'cilic  'j:ia\ily  liy  tlir  cnii- 
■t.iiit   iMicflicicnt   0.'_':i:l  I  Ilai's(>r'>. 


^>       f 


The  Depression  of  Freezing  Point 

NVIiili'  till'  soliils  111'  till'  liliMil  I'liiisist.  I'm  ihi.  must  |iait,  ul'  pripti'iiis 
iiiil  ciillniils.  Ilii'si'  III'  tlic  iiiiiii'  an-  nniili'  up  ul'  iihnsraiiii*  salts  ninl  small 
■  •r._Miiic  iiioli'i'iili's.  Till'  iniili'riil.ir  i-nni'i'iitratiiin  tliat  is,  tin-  total  miiiilii'r 
i.r  iiiiili'riili's  in  a  yivrn  ipiantity  nl"  tliiiil  is  nmli'r  orilinary  I'onilitioiis 
iniirli  irrrati-r  in  tin-  iiiini'  tlian  in  tin'  lilooil.  Tin-  nioliTiilar  I'liiu'iMitra 
;i..ii  ni.iy  Ii(>  ili'tiTUiini'il  liy  tlie  ili'prpssinn  oi"  tin-  fn'r/iiii;  ]ioint  of  a  fliiiil 

'  '  I'u    tlKit   i.f  liistiJli'l   wati'i'    I  ^.'i'   pi'ji'    I'll.     r,l 1    j'ri'i'/i's  almost   •■mi- 

-iiiiitly  at  O.'il  ('..  wliili'  iiriiii'  iiia>-  fii'r/r  at  \  arinlimis  of  tompiM'atiifo 
'-.■iwi'i'ii  1  ('.  aiiil  '2.'>  < '. ;  if  M'ly  i-oni'i'iitiati'il  it  may  fri'ivo  at  a 
'.  iiipi'ratiirp  as  low  as  ."i  (".,  or  if  ililiitr  tin'  rriiv.iiij;  point  may  lip  as 
i  idi  as    0.07'."  C. 

Tlip  v;:ii;ilpilit.\-  of  tin-   fri'iziii'.'  poini    ami   ll.i'  sj>i'iM(n'   uravity  of  tlio 

i'iiii'  loail  us  to  a  i-oiisiiliTation  of  tin'  fi'latioiislii|)  of  tlio  urinary  vohimo 

I'p  its  I'oiici'iitiatinii.     In  tlio  first  iilai'i",  tlie  voliimi"  of  \\ati'r  iniri'stcil  is 

'li.iii'  fri'(|ui'iitly  tliaii  otliiTwisi'  in  I'xi'i'ss  of  tlio  miiiimuni  alisolutoly  vo- 

uirril  hy  tin'   lioil>'.  ami   is  suli.ii'i't    to   irii'atiT  variation   tliau   tlic  sub- 

■iitii'os  oxci'i'tril  ill  llii'  iirini'.     Tlu'  kiilnoy  is  alilo  to  oliminato  one  pon- 

'itiipuf  of  till'  iilaMii,-!  wliii-li  may  lio  pri'sont  in  pxi-r'ss  witl'out  itivolvitiir 

:i\  i'lian>.'i's  in  otln  is.     I'or  pxamplo.  wlim  salt  is  aiMi'il  to  the  fooil  anil 

M'i'i'ti'il  in  till-  uiini',  tlip  total  I'liloriili's  aro  iiii'ii'asi'il,  Imt  tlio  amount 

''  iirino  ami  tlio  otlior  oonslituouts  may  romaiii  uni'li.iiiiti'il :  or,  at'ain,  as 

:iy  liaiipon.  oxooss  of  salt    loads  to  an   im-roaso   in   tlio   volume  of  the 

nn-.    lull    ilio    sail    oonooiil  ralloii    lomaiiis    ooiislaiil    wliilo    lliat    of    llio 

lior  tiriiKirv  lioilios  is  ilooroasoil.     Siniilarlv.  altliou"'li  tlio  rate  of  \irea 


! 


I 


I!  I 


'i 


\i  - 


u\^ 


I 


Hi:i 


4h 


!i 


I  III      I  \i  III   llliN    liK    I  HIM 


I'XiM'cliuli    is   lint    iliiiiiilis1l;ilil,\    iilliriili'litiil    li\    ,'ili    iiii'ii>;i-c    ill    tlic    viiliin 
111'  tlic    lirilic,   ;iM    ilKTi'iisr    ill    til,.    I'iilc   111'    iiiiM    i\i-|-ctiiili    llicliiccd    liy    (!■ 
illiri'stiMli  III'  lllrn   i>  !li-fnill|iilllii'(i   li\    ii   1,11  Ufl-  \iiluillf  i<\'  lllilii'       'I'llflt   till  - 
twii   t'iicliiis  Mi;i\    iiiit   st.niil   ill  ,'i   (•;ii|siil    ii'ljit  i>il|Niri|)  U>  imcIi  utlicr  is  siiv 
irrsti'd  li.\    ii'irnt  \Miil<  lit'  Aijilis  ,iiiil   Wiit.-ilKilic,    who  I'liiil  III!  i|iiiiiititiiti\ ' 

ri'Illti()lls||i|)     liclWclM     till'      IMlc     III'     illrll'.'isc      ill      lll'cil      I'XlMclinn      tlllll     l|i 

iiicfc.-isc  ill  iii'iiiii  vcijiiiiic,  iiiiii  u  III.  Iii'lii'\i'  tlijit  llii'  ii|>iiiiii'iil  ii'l;iti<iiisliii. 
is  line  til  M  I'liiiiiiiiili  I'liilsf.  siH-li  Ms  .iltriMliiiii  in  llic  liiti'  nt'  I'i iciiliit iiiii  "• 
clllllll.'''  ill  llir  iii-ti\  it  V  III'  ihr  l.iillii'V  it  I  Is  \r\  cillirlrss,  tliiTi-  ;i|i|m'1IIs  I- 
III'  a  limit  si't  til  llii'  |iii«i'f  111'  till'  killing    in  tjilii'  llir  iiiiiiiii\   sjilts  m-  w^it.  i 

frnni    ilii'   |>l;islMil   allil    t   i   |ilili'r   tlll'lll    ill    tllr   lllinr   ill   i|llitl'  ililVrli'llI    |ir.i|l.' 
lions      Till'  ili'liiiiti'   Miiiiiiliit    111'   \\.itii'   ii'i|uiri  i|   in   Imlil    tlir   iiliiiiil'v    s,il|, 

lias  liifii   triiiiiMJ   till'  "viiliii lilii.'aliv  .•■'      Aiiiliaril  Tin'  i'    limits  :,• 

roiii'i'iitratiiiii  iiiav  In'  liNfii  li\  tlir  nii  iltx  «  liirli  llir  Miliicy  i-aii  liriim  ' 
ai-f   airaiiist    tin'  nsiiintir   i  I'sistaiiri' 

'I'lii-  iiirniistanr.v  ill  till'  lii-lia\  iiir  nl'  till'  kiiliiry  luwaril  iii'ji'sti'il  siills  i. 
in-iilialilv  iliir  til  till'  I'ai't  iIkiI  iIh'  sjiIk  umiIi  tlir  kiiliii'v  in  tin-  niiii'i  i 
tiatiiiii  ill  vvliii'li  tln'\  aii'  lii'M  li\  tlir  liliiml  plasma,  ;iiiil  nut  .'is  tlii'v  "•■- 
iiitrrstiiil  I  r  s.ili  is  alisiiilii'il  i;i|iii||\  I'linn'jli  til  ilisinrli  tin'  salt  i'i|iiiliii 
liniii  111'  till'  lissiii's  ;iiii|  |ilasnia.  tlii'ii  \\,ili-r  «ill  ln'  alislfai'tnl  IVinii  tl- 
tissiii'v  aiiil  till'  plasma  uti  irarliiiit!  llir  Uiilni'v  x\ill  iliminati'  tlii>  s,i- 
ami  >  ili'i-  liiLirtlii'iv  Till'  iJilTi'irri'i'  in  lli.'  rfai-linn  arises  I'liiin  ll  . 
viii-ii'il  .-ii'tivitx    in  till'  tissues  in  L'riii'ia!   ratliri-  tli.in   in  tin'  kiilni'v    itsr!' 

The  Reaction  of  Urine 

III  man  ami  tin'  rainiMira   lliis  ifai'limi   is  m'lii'iall.x    ;ii-ii|  to  litmus  n^ 
plii'iinlplitlialiin.     Till'  cause  is  riMiinJ    in  tlir   I'iii-t   tlnit   llir  rinl    pniilnct 
iif  piiiti'in    mrt:iliiilism    yivr    risr   tn   siilplnuii-   .-iihI    pliiispliiii'ir   ariils   tli 
ariility   ut'  wliirli   uivrs  llir   niiiir  im   .iriil    irai'limi.      In    tlir   lirrliiviiruii  ■ 
.•liiiinals    till'    alkaliiir    ir.'n'tiini    is    ilnr    In    tlir    I'.ii'l     tliat     \  riirtjililrs    an- 
fruits  r.intaili  salts  nf  ilili,isir  m    piil\  lijisir  ariiN.  siirli   as  ariil  piitassim 

iiial.'itr.   ritratr,   arrtatr,    anil    tai  tijitr.      (Kiilali !'   tllrsr    in    till'    liml 

•ii\('s  risr  tn  r.'llliiiliatrs.  Snmr  iil'  tlir  railinllir  ariil  is  rxri'rtril  tlllnU'.' 
till'  liiiiu"^,  ami  lirnrr  tlir  a-  ■  irial n  1  liasr,  Lrnirrally  sinliuiii  111'  iiotassinii 
is  riiiiiliiiiril  sii  as  111  rnrm  h  wrjik  liasir  salt. 

Tlir  iiii'.Msni-riiiriit  111'  till'  ariilit\  <>(  Ilir  niiiir  ill  tiTnis  111'  irr.'im  aiii"! 
Ill'  ratiiills,  likr  till'  samr  liirasii  iriiirnt  in  liliiiiil,  rripiiirs  tlir  nsr  iif  tl, 
ratlin-  iliflinilt  rlrrtriral  m-  imlir.ilnr  mrtliuil,  tlir  piiiiriplr  of  w  liirli  li.-! 
Iirrn  ilrsrrilirti  in  <|i.iptrr  \'.  I'I  \  tilrssri  I  in  t"l'llls  iif  (',,.  tlir  ariili' 
wii'ii's  lirlvM'i'ii  4.7  \  in  "  aiui  lilii  \  III  ■.  ■/■/(,  iiiliil  iiiilnilidl  udditit 
tliat    is,   till'   lilimlirr  :>(   ||    imis   uliirli    ivill   lir    I'olliU'il   ill   tlic   fare  nf  a   >■>■ 


\Mor\T.    (  (i\llii>lllii\.     \Mi    f  II  \ll  M    11  It    i'|-    I  IIIVI 


.r_'.i 


lihilJil    Mi'lllnili/illicili    III'   lliiisr    in    ^iilutinn      lilllN    lir   iili(;iil|ri|    I'iiiily   iicrii 

!;ilcl\  li\  tiliiiliii'.'  tlif  iiiiiic  \\\\]\  ',,,  liiiiiiiiil  iilkiili  ill  till'  in'i'sciiec  ol' 
iiiiltnil  iHiliisviiiiu  nNahiliv  ii^ini.'  |ilii'iiiil|i|itliiili'iii  :i>  iin  iiiilii-iituf  (FolinV 
Tlic  ifslllls  \ir.\\  III'  i\|iri'ssril  ill  aciility  prr  rnil  ill  ti'lliis  nl'  cc.  \  Id 
N.lOU  ifijlliiiil  111  IK  llfrali/i'  llM»  !•  c-  nf  lililHv  If  tlir  iiliiliinliiji  i-Xrl'ct  iiil) 
i.  .iilili'il  111  till'  lili.iliiiii  i-i'MiIIn,  till'  tntiil  |iiitriiti:il  ai'iility  is  mtv  clnsclx 
iiiiiisiiri'il. 

Till'   iiiiiii'    is    iiiiiic   alLiiliiii'   sliiiitly   iil'lrr   iiii'iiK   llian    at    iitliiT   tiiiirs. 
.iiiiT  ai'iil  is  liriiii_'  c  \cri  lid  li>    till'  i.Msliir  ijlamls.      It  is  iiidic  arid  on  ii 

al    than   mi    a    \  I'U'i'laiili'   dirt,    and    is   arid   diliiiiu'   stnfvation    lu'i'iuisi' 

IHiiti'iii  is  tlit'ii  tlir  idiiil'  iiictal'iiliti'.  Ill  discasi'  llii'ii'  is  nn  idiaiai-tffistif 
\aiiatiiiii,  sa\f  lli.il  tlir  iiiiiii'  is  wrnvr  miiciallv  acid,  wliirli  iiiav  lif  cn- 
|ilailird   li>    till'    tact    liial    in   siiimis   iliinss  tlic  dirt    is   list  lictrd.      When 

till'  ai'idit>    is   iiii'irasfd,   il MTi'limi   nl"  aiiiiiiiiiiia    is   iisuali\-    i;ri'af<'i'. 

sJiiiM'  aiiiiMiiniiiiii  railiiinati'.  tin-  rmi'i  iiniH'r  nl'  urea,  ads  as  an  alkiili  and 
iiiiitrali/rs  till'  aiid  ladiidi'-  'I'liis  i  isc  in  aiiiiiinniii.  lii>\w\  cr,  is  iint 
:il\\a\s  |irii|iiii  liunal  In  iIh'  arid  ladirlis  prisint,  since  tlic  li\cd  alkali 
i|i'ii\i'd    rriiiii    iViiils   and    \  c^dalilcs   may    In'   siil'licii'iil    In    nciiliali/c    tin' 


I'id    full 1. 


THE  SOLID  CONSTITUENTS 


i'l.i'  jiiactical  icasiiiis  w,-  shall  di\  idc  tin nstitiicnls  nl'  tlic  urine  info 

hiiiiiial  and  almiii'inal.  'I'lic  rniiinr  arc  picsi'iit  in  the  a\cras,'c  nrinc  in 
:iiii(iiinls  s\il1iciciit  til  lie  ditcclcd  liy  (irdinary  means;  the  latter  f)lily 
laicly  appear  in  detectalilc  ipiaiit  ilics.      )ii  a  person  eatinu'  an  ordinary 

diet  tlic  most   iiiipnitanl   nruaiiir  and  iiioruani mstiliients  of  tile  tirine 

all'  as   follows: 

TiiiAi.  Sol. ins    I  III    Ml  C'li  Ciiwis.    i\   (IM    I.I  in:  hi-    Ni.i;m  m.   l"i:i\r 


"ICI.ANIr    |ii.\.s|  I  1  11  N  I .-..    I'.'i    III   i:m. 
I   It  :i.  I'll  ;'.."i  ^111. 
I  II  Mliiiini'.  l.llll.."i  ^ii\. 

mill,  II..')  l.J.'i  ;;iii. 

Ili|>|iuiii'  nciil.  0.1   1.7   :;iii. 
iiiliiT     riinslitiirnls     1  illii'i.iil     sMlpIiit. -.. 
'•\:ilii*   Mriil,   iiiiii:t'\    j.i-iin'it^.  'If.  >. 
I..'i  l2..'   L;in. 


i\(ii:i.A\li'  eossTii'ii  N  IS.   I."i  :.'.'i  i.M. 

S.i.liuill   illlnli.li'    I  .\';i('l  I.   S-I.'l  KIM. 

I'll. is|ili, nil-  ai'i.l    I  l'().  1.  J.."  ."...")  yiii. 
Siil|i|iiiiic'   iii'i'l.    (."sO,).   '_' L'..'   yiii. 
I'.iiMs^iuiii   I  K.i)  1,  :;  .'!  yiii. 
S.i.liuill    1  Nil  (» I.    I  Ii  yiii. 
<':iliiillii    I  CmUi.    (1.1  II. .'1    i;lll. 
M:i:;iii'simii    (Mi;()K  0.-'  Ii.."i. 
ViiUii..iii:i    (  Nil   >,  11.:;  l.'J  u.n\. 
Ii..ii    I  ill    i.iyii'.rnf',  (i. full  (I. din. 


r,,iii|.il..l   Iri'iii    \lii-iiitli;il  '-I  Ii::.!!.  -. ' 

Tlicse  urinary  sails  ai'c  pn'scnt  in  llie  lilood,  and  are  excreted  only  liy 
he  kidney.  An  iin  (■stiyalioii  of  the  mechanism  of  renal  secretion  must 
hcrefore  inclinlc  a  sinii\    if  liie  rciatioiiship  cxisiiii!.'-  lietwcen   the  coii- 


i| 


■J 


h 

I'r 
!  ,, 


H     ii 
if.;  11 


!-ll 


Iralion  of  the  niiinirv   salts  in  the  hi 1  and  in  the  urine. 


iXla 


hi,  t 


■  >-i>  Till:  I  \i  III  Tii)\  iiK  I  him: 

The  Normal  Organic  Balti  of  the  Urine 

Nitrogenous  Constituents.    Tlic  u'lmi.  r  ihiiiiIm'i'  of  t|i,>  niirniiic  s.ilu 

till'  iiiiiir  ill.'  iii.i.li'  ii|>  uT  liiiclio  uhidi  ( niin  iiitii(t;,.|i,  Mini  wlijc'h  ,■, 

i!('ri\'il  I'lurii  llir  pruti  in  clrni.iit  of  iiiitriticin.     Tlii'  |H(itriim,  \\U'i,-]t  fi,!.,, 
till'  chii'f  liiiililliii.'  iiiiilrtijil  of  till'  lioily,  .III'   liroki  11   ii|)  into  tlicir  <■'■ 

xtitlli'llt    iinillio   iiiiiK    ill    tl;r    iliti'sliiiiil    ti'ii.'t    mill    III  s..||,,.|l    iis    siK'li    la    tl 
liloiMJ.      I'oilions  of  iIh'^"  ,11'iils  Jill'   liikcii    up  |p,     thr   tisMH's  |o   H'|(iiir  ill    1 
to  icpliH'i'  llioM'  piotiiiis  wjiicli  liiiM'  lii'cii  iJiM'iiidi'd.  ami  tin'  ii'iiiiiiiiin  ' 
proti'iii.    ill   I'M'.sv   i,f   till'  I)  mI.\    III  I'll    f,.i'  iiiiiiii,!   iii'iils.    is  ili'jiiiiiili/i'il.   I' 

in.'i.jor  portion   of  i| iiilinn.   o\y._'i'ii   ami   liv.Iiovcii    liciiiir  oxiiji/i'i!    '  ■ 

foiin  ro    jHiil  watiT.  anil  tlic  Ii'smt  porti f  tlii'M'  ili'iii.'iits  licini:  I'on 

liincil  with  till'  nitioL'i'ii  to  forni  iirca.  iiiiiiiionin,  m-i,.  arid,  etc,     A  siinil   ■ 

fiiti'  lati'i-  awaits  tlu'  nit  lo'.'i'n  iiioi.'tv  wliii-h  foiiiuj  a  pli in  tin-  tissii, 

.'Hill  whii'li  is  ii'plai-iil  in  tiini  li\    nt'«   nitro'.'i'iiMiis  lio.lirs.* 

Siiii'c  all   till'   inu'isti'il   nitroi_'.'ii,   t  xccpt    a   small   ami    tatlii'i'  roiistji    i 
aiiioiiiit  wl.ii-li  is  lost  liy  ill.'  fi'i'is  ami  llii'  swrat.  is  cxiTi't.'il  in  tin-  mil  ■ 
llic  total  nitioiriii  of  till'  iiiinc  lias  lu'vii  taki'ii  as  a  iiii'asiii-i'  of  the  niti 
','i'M  or  pi'oti'in  iiii'taliolisiii  of  tin'  limlv.     hi  iiorinal  I'oi  ilitiuns  tlic  prot'  i 
iiii'tal.olisiii    is   jmIJiisIi'iI    ill    siii-li    a    iiianmr   that    the    iiitrojri'ii    iiit.-iKi'    i> 

I'lpial  to  till'  iiitroiii'ii  output,  ,- mlilioii   Known  as  nit  ro'.'inoiis  I'ipiili' 

rium.      If  till'  nitro'.Tii   intaki'   is  ii'iIihtiI   lidow    tlic  actual   limlv    miij 

'• M'rrtii.n   of  iiifi-.>-r:i    is   ..'IV  Iter   lliaii    I'lc   inl.iki'    ivliicli    in  liciitcs   lli.  • 

the  lioily  protein  is  II  placim:  the  protein  Usiuillv    fiirnislieil  l,y  the   foo.i 
The   niininiuin  amount   of  protein  tli;it   the  Imdy   niiist   lia\c  to  inainl.r 
ei(iiililiriiiiii  x.irii's  in  imli\  idiials.  Imt   is  on  the  averauv  lutueeii  Ti  and  >■ 
'-'laiiis  of  iiitioeeii  a  day.  wliicli  corresponds  to  alioiit   ^O  u'raiiis  of  pi 
tein.     With   the  ordinary  diet   it   is  iiMially  Itetwceii    12  and  2(1  uraiiis 
day,  or  represents  from  7"  to  12.">  ■jr.iiiis  of  protein.     .Since  protein  js  le  ■ 
stored  liy  the  liody  cM'ept  i  i  periods  of  ci  ou  th  or  after  periods  of  iind 
nutrition,  an  increase  in  the  protein  t'ood  is  ;iecoiiipaiiiei|  Iiy  an  iiicrci 
ill  the  iiitrie.'eii  e\.'reted  in  the  urine,     j-'or  this  re.isoii.  unless  the  atiioU' 
of  nift'oyeli   ilit;estei|  is  klioi    11,  the  stild.v    of  the  total   nitnej^en  of  the  111  i: 

U'ives  no  iiifm mation  eoiiee  nini.'  the  nature  of  the  nitio<ren  metaholi^ 
of  the  liody.  The  total  output  of  nitrou-en  per  day  usually  amounts  • 
111  to   |.">  yr.tiiis      fn. Ill    1    to  2  |ier  cent   of  the  urine  liy  wei'.dit. 

All  the  nitrie.i-e s  Imdies  of  the  urine  arc  nornially  imiiprotcin.  ai    ' 

arise    from    similar   liodies   in    the   hlood,    m  here   they   exist    in    cm nli 

tioiis  of  from  20  ',1  :;(l  mir.  per  tun  ,..c.  In  exereliii'.'  tlie  iiitr()«.'cii  of  ll 
urine  the  kidmy  therefore  takes  it  from  a  solution  in  which  it  is  foiii 
'">  "  '■<'! nti-aiion  of  ii.d:',  p,.r  ....nt  on  the  averau'e  and  delivers  it  to 


'  l''..r  I  iilln  I  .!,  I.,  1..  ,,  V  p.i«r  f,ln. 


AMIIINT.    I  (IMI'IINITIOV.    AMI   (  llAKXrTIK   IIK    I  HIM 


~<>liitii)ii   i'<iiit!iitiiii^  an   ini'in^'i'   nf   I  ittl,   ni'  ciiiici-iitiiitcs   it    at    It'ast   -tO 

1  lllll'S. 

Urea.     Tl lii<-l'  nf  tlic  iiiti'iimniiiis  limlii's  oi'  tlir  uiiin'  is  iitra,  th** 

Ki  lU'iil  ••!'  wliii'li  has  liccii  l'iill\  iIi'sitIIii'I  in  tin*  i-liaptirs  i>ii  inctalMiliHrri. 
Nil  ronstitiiciit  nl'  tlic  iiiiiii'  ]•*  siili.i"'i'l  tn  iri"'ali'r  \aiiatii>ii  Imtli  in  aliso- 
hilr  anil  in  I'l'lativc  ainiinnl>.  On  un  mm'Iml;!'  dii  t  mhiiiinini;  120  (trains 
III'  |iiiili"in  |>i'r  ilay.  tlir  iilisulnlc  uira  cMTi'tiiiH  may  aiin'iint  tu  .ilimit  ISO 
■.'laiiis;  (111  a  linv  iMiitcin  ilirt  it  may  In'  only  a  I'm  i;iarMS,  Wlnii  tlic  |irii 
t<  ill  iiitaki*  is  liiuli,  tlic  iiilruL'cn  cliiniiiati  il  as  iirca  iiia\'  lie  !HI  pci'  cent 
il'  tlio  total  iiilniircii;  lint  wlicii  tin-  prntciii  intake  is  jnw,  tliis  pi(i|mrliciii 
iiay  fall  t"  tHI  pec  cent.  'I'lic  ililVercncc  is  lieeailse  1)1)  a  low  Jimteiii  ijie! 
till'  tricatcr  pei'ecntai.'e  nl'  iiitnii.'en  eliniin.itcil  is  cniliiu'ciiuiis  in  nrinin, 
.iinl  iiiea,  \\  liieli  is  tlic  eliicl'  ciiiistitneiit  of  tlic  cxiiirciKiiis  nilmifi  n  iiiiiiety 
III    tlic  urine,  is  aceniilinirl.N    (jeclcascd   mi   Inw    diets. 

In   recent    ,\ears  llie   impnrtanee  nl'  llic   rclaliiiii>liip   lictueen    the   cim- 

iitratiiin   nl'  the    iiinaiy   cnnstitneiits   in    tlie   IiIimmI   and   llie   urine   lias 

I' ell    niilcli    iiisis'ed    lipeii,    aiid    since    (lie    I  >l  Iniation    nl'   tlic    ainiiinit    ol' 

Miea   in  tlic  liliii  d  and  lie  nriiic  is  relatively  sniplc.   ninsi   nj'  ||ie  wmk 

\\;is  liecii  dune  liy  iisiiiir  tliisc  values.     Aniliaiil  and   Weil     li'lievc  that   a 

ititati\e  rclatimisliip  cxInIs  liel\\i''n  tlic  rate  nl'  niine  cxiMctinn  and 


•  pii'ii 

ihe  eiinccntiatioii  III'  urea   in  the  Mnnd  nnd   tlie  iiiiin 


the  liliinil  acts  as  a  stiiniilns  to  tlic  icnal  ei 


H 


since  the  urea  in 
V  eniiipariii'.'  the  rate 


III'  urea  exerctinn  and  the  enneentratinn  nl'  urea  in  the  lihmd  and  urine 
in  a  inatheiiiatical  rnriniila,  they  li:ive  nlitaincd  a  \aliie  whieli  they  lie 
lievc  is  'aiire  or  less  lixed  t'nr  the  nnrnial  kidney.  This  e\|iiessinii  is 
I  iiowii  as  Aiiihiinl's  rm  niciiiit  iiinl  fdniniln,*  and  has  lieen  used  as  a 
•iie.'ins  of  evaliiatini,'  the  functional  capaeitv    of  the  kidney. 

WhalcMT  the  value  of  the  tiirniiila  iiia\'  Ic  in  expressiii<_'  the  relationship 
I  sistinc  lictu  ecu  the  raie  nf  iir.'ii  cNcretinn  jind  the  enneentralinn  of  this 
-lit  ill  the  liliind,  it  is  certain  that,  in  diseased  eniiditiniis  xvheic  inip.iir- 
iiieiit  of  tlie  kidiie\-  is  ceitain,  the  eniieeiit  r.il  ion  nf  iirea  in  the  lilnnd  re- 


liiis  iieriiianeiitly   at   an   alinniinally   hiirl 


I   aveiaire 


level,    althnllu'h    tl 


■.\?iili;ir.I    .-jTi.l    W'l  M's    f..riliul,i    U 


.■iMffiiiir.t    „f 
K'Jinf*   III    nri  ;l   j  i  r 


Ir 


lil.r  nf   M".'.l-. 
iitiMit   of"   nrrn    ill   cr.iiiiv    |.i  r    '4 
e  »rlKllt    111    the    I  .Iti.  I  1 

t"  «r;itTis  i.f  Mif.n  itr  titi  i    ..f  uniic. 

7(1         sMii.l.inl    wriuht. 


ni       \  »• 


ll'..-:-t^l!  I    .  f    .\!mI  ;,r.ll 


.«t;iiii]iiril   I  iiTiitMtr.itii 


■III.-    .IV, 


alii.'   fiii    III! 


lltWi.lMt       III      tlr.MI. 


I  riliral    rr\ii«.    nl    llir    »irk    li.i\r    In  ill     |iul.li-!ii  .1    nriiillv    liy     .Marl,  .in''    ami     l.y     .Vil.li- 


i 


;^ijj 


Mil 


n 


li'^ir 


M'.' 


.< 


:!il     .    » 


liii 


Hi 


Iff 

till: 


•  il-'S 


I'm     I  \i  l.'l  IICPV    n\-    I  l;i\i; 


.■miipiiiit  III'  iiri'ji  cxiM-rlfil  iliiriiiL;  Iwiiilv  roiir  liiiurs  iiiiiy  he  cxactlN  tin 
sMiiic  iis  iiikIiT  inu'liiiil  cciinliticiiis.  I'nilitililv  llic  iiici'i'iisi'tl  <-iiilct'liti  iitim 
'>\'  Ufi'M  ill  till'  lil(i(p(l  uiiiltT  llicM'  ciiiiililiiiiis  'k  ;i  ciiiiipciisMlciry  iiM':isii!> 
111  pniviili'  siil'ticii'iit  prcssiirt'  tn  riiiiM'  ils  cxiMi'tinn  tlifmiu'li  .'i  iliiiii.-iiiol 
niiili't.  It  i^  lliis  iiici<>asc  ill  iiii'ii  „\'  ilic  lilodij  uliidi  is  imliciitt'd  \>\  tli^ 
Iciiii   iiriii  ntiiilidii  ill  ticpliiitis. 

It  must  licit  111'  Iiist  sii;lit  III',  lui\\c\cr.  tliiit  tlic  appnixiinato  coiistain 
111'  the  (•(iiiiliiiicd  t'liinnila  is  clue  in  hiri/c  part  tci  tlic  iiiatiifiiiatical  i-ni, 
slniriiiiii.  and  aUci  tn  llic  fai-l  that  ari\  iiiiT.  .isc  in  tlic  I'lincfntratinii  ><( 
urea  in  tlic  lilcicid  is  iisuallv  acciunpanicd  liy  an  increased  latc  of  iiii  .1 
excrcticin.  The  lactcus  whieli  are  iinist  variaKle  ciccnr  as  tlic  s(|iiai'e  ..' 
the  sc|iiare  rciiits  (if  their  \ah|es,  ami  thus  the  distnrliinci'  ciTcct  tliey  pi.. 
dilcc  cm  the  ciiiistaiiex  cil'  the  re-uhaiit  cit'  tin.  rcniinila  is  irrcatly  1. 
dilc-ccl,  while  the  must  ccnistant  lactm-.  the  ecineeiitratinii  nf  urea  in  ll,. 
Iiliicid,  is  used  with  iiicidilieatidn.  In  such  a  cciinplcx  niec-hanisni  as  th. 
renal  I'lnicticiii  it  is  xcrv  prciliahlc  that  cither  t'aet'irs  are  iij'  jji-eat  iin 
p<irtanc-c  in  cuntrcilliiiii  the  rate  cil'  urinary  excretion.  Many  nj'  tliesi 
t'ac-t'irs  can  ncit  admit  cil'  mathematical  cx|>ressiciii.  The  writer  scriciusl\ 
ihiiilits  the  acl\  isaliility  nf  adciptinir  an  empirical  fcuiimla  as  a  iiieai,- 
iil'  exprcssincj  iiiikiiciwn  pliysidlin^ie  laws.  Such  measures  are  apt  i. 
U'ive  seii>e  cij'  kiMiwIednc'  altte.'cl  her  false,  aiicl  thus  hinder  rcseaiil 
pr<ii.'rcNS. 

The  upper  limit  cit'  McmmI  urea-nit  rciu'cn  is  ahcnit  "JO  iml'.  iier  101)  |.  ,• 
which  wiililcl  ciirrcspuiicl  In  aliciilt  (I.4-")  ■.'iii.  cit'  urea  per  liter  of  lilm.il 
The  avcrai.'-c  li.iiurc  is  half  of  liiis  anmunt.  Tiic  maximum  eciiicentratii.! 
of  urea  in  tlii'  urine  is  seldom  over  S  per  cent.  On  this  liasis  the  kidnc  \ 
c-aii  raise  the  ciuieeiit  ration  of  the  urea  in  the  iiriiic\  at  a  ecin>cr\ati\  ■ 
estimate,  from  10(1  to  'Jotl  times.  Ndrmally  the  daily  output  of  iire.i 
nitroifcii  iii.iy  ij.iil'c  froni  S  to  12  'j.\\\.,  and  the  nitrocjen  wliicli  if  contaii 
is  rciiiLddy  .^0  per-  cent  of  the  total  excretioii   for  the  day. 

Ammonia.     The  cdiicf  >ciiircc  of  ammonia   in  the  hody  is  from  the  ni 
t  idsreiious  poitiiiii  of  the  dcaiiiidi/ed  amino  acids.      The  aiiiiiionia    foiin.! 
in  excess  in   the  porta!    Iilood   is  derixcd    from    iiiiicstcd   ammonium   s;il;- 
aiicl    from    amiiionia    resiiltiii!,'    from    luicterial    action    on    proteins    in    t! 
intestinal  tr.ict.     The  ammonia  of  the  Imclv    is  present  cliietly  in  the  fori 
of  ammonium  carlionatc,  ;jiicl  i'   is  this  salt  that   is  the  precursor  of  uic 
liccaiise  ammonium   earlionate   is  so   reacjily   converted    into   urea   hv  tl 
tissues  of  the   Imdv  ,   little   alilliiciuia    is   norinally    present    in   the   systeiii 
liloocl.     The  irrc.-itcr  portion  of  the  ammmiia  that   finds  its  way   into  tl 
urine  serv,  .  as  a  hase  t.i  transfer  acid  radicles  either  iiiu'csted  or  fortie  ■ 
within  the  liody.     The  amount   of  amiiicHiia   in  the  urine,  therefcu'c.  is  ;i' 
iiiclircct   measure  of  the  CAteiit  of  urea   t'oriiiat imi  ami  of  the  acicl  Imdii 


\Miir\r.  ( (i\ii'(i-i-nciv.    wn  im  \i{  mti  is  in'  i  ifivi: 


."2!t 


,,r  llic  liliioil.  l''i>r  llic  \;\\Ur  iciisciii  llic  (li'ti'|-inili.-i1  ion  of  llic  iiimuoiiia 
.  sclftioil     ill    lllilll'    is    of    suliH'    I'iillicill     illllioit;il 'rill"     illf^fsliou    of 

iiiiiifriil  iU'ids  iiicicascs  llio  iiiniiioiii:i  cxcii'tioii.  wliili'  ;illi;ilit's  tend  to 
rcilnco  it.  Diii-iiiK  fiistiiiir  t\\u\  in  ilisc;i>fs  Midi  iis  diiilictcs,  wiicrc  tlifii' 
i-;  an  iiliiioi'inal  iiirtaliolisiii,  lln-  aiiioinit   of  aininohia   in   tin-  urine  is  in 

i-rcasi'd.      <  Miliiiaiilv    tin-   daii\'    ont|inl    of    nun ia    nilroij-cii    docs    not 

,'\i'i'cd  0.r)(t.(i  •iiri..  coiistilntini:'  ;'.■'>  \<rr  i-i'nt  of  tlic  Iota!  aiiioiuit  oi 
iiitrosit'ii. 

Creatinine.     <>n   a  meat  frci'  dii't   tlic  dail>    i'\i-i-t'tion   of  iTfaliniiii'  is 
icniarkaMy  constaiil.  ainoiinliiiL'  to   from  7   to    II    iiii--.   pi'r  kilo^'rain   of 
l.ody    \\t'i!.'lit,      l-"or  this   ri'ascni    its  dcli'iiiiinalioii    is   ai'i-rpti'd    as   an    in- 
■  lispciisalilc    tVatiiif    ill    nictalioiisiii    invcstiuMtioiis    iiivniviii'.'    iiriin'    an 
al.vsis. 

Any  iiross  \arialion  from  tlic  noimal  ainoiinl  iinlicitcs  tlit>  rfrtain 
failui-c  of  till'  atl.'iidaiits  to  colliTt  all  of  lli.'  t  u  i'iit\ -four  Imiir  s|ifcimcii 
of  \    iiit'.     Normally  tlic  hlood  conlains  from   1   to  l  iiiu-.  jicr  100  c.i-. 

Tlio  fi'catiniiu'  is  one  of  tlif  last  of  the  nriiiaiy  coiistiUK'nts  to  ai-i'iiiiiii- 
hito  ill  tlic  lilood  duiinu   ii'iial   iiisnt'licifiicy.  and    for  lliis   reason   atl'ords 

,-i   rt'liatili'   )irosrnosti('   indication   co rniiin   tlic    ]iaticnts'   <'oiiditi()n.     A 

II-'  ill  the  creatinine  coiii'cntratioii  of  tlic  hlood  is  c\  id.'iice  of  serious 
rciial    disease,    (latieiils   with    concent  rat  ioiK   of  .">   iiii;.    ncsce   reeoveriiiii 

('liase  and   .Me\crs  ■     Tli mccntratioii   of  creatinine   in   tlie  urine   is 

.iliiiilt   100  tiiiii's  i.'rcatcr  lliaii  in  tlie  Mood. 

I  I  adult  man  creatine  ilofs  not  anpear  in  tlic  urine  save  diirinu-  starva- 
lioii  (H'  wastinii  diseases.  In  woman  it  is  aliseiit  sa\c  after  postpartum 
resolution  of  the  iitcnis.  ('hildrcn  conimonly  excrete  i-reatine  ahniu' 
vith  creatinine  until   the  middle  years  of  childhood. 

The  Purine  Bodies  and  Uric  Acid.     The   nmst    imporiant    iniriiie   in 

iiiniian  urine  is  uric  acid.     Xanthine  is  the  next   in  iiiiportanci'.  and  small 

.iliiollllts  of  hypoxanthilic,  i^naninc,  and   adenine  are   found.      Aliionir  the 

most  ilitcrcstinu  of  the  salts  of  the  urine  In  the  eliniciaii  arc  the  urates. 

Iiecaiise  an  acciimiilatinn   of  uric  aciil    in   the   hod\    \>as  hclicvcd   to  he 

icsponsiblo  for  many  ohseiire  clinical  conditions.      It   is  rpiite  true  that 

•lie  salts  of  iiri<'  acid  are   fmiiid  in  hiuher  than   noriiial   amount    in   some 

diseases,  especially  iroiit.  leukemia,  and  (dironic  nephritis,  hut   the  iiian> 

airue  theories  associated  with  uric  acid  aiel  disease  h,i\e  lon<r  aiio  Immmi 

xploded. 

The  liiiiiian  liod\    has  the  almost    imiiine  distiiu-tion   jiiiioin.'  mammals 

f  not  heiuii  aide  to  destroy    ali.\    of  the  uric  acid   it   iirodilees.  and  hein-c 

ill  the  uric  acid  t'cn-mcd  durinii-  inetaholism  must  he  excreted  in  the  urine 

!!ifortunate|y  the  kidney   .-ippea.rs  t(!  he  h'ss  .•omjuMenl    to   lid   the  lio.ly 

'<(  ihis  waste  than  it   is  of  the  other  urinarv    metaholites,  and  one  of  the 


it! 


tl 


1 1{ 


iii-  ^ 


i    I 


•'■■■  r^'1 


I 


■  .1     i 


I'  tl'  '1 


MO 


Tin:  i:x(iii:Ti(i\  ok  ikim: 


.•iiilicst   siiriis  of  ivMj.l   iiisunici.'iicy   is  ii<i\v  lirld  to  lie  a   failure  of  tl;. 

lu.liicy  f.,  pivviii   tiic  iiiic  aci.l  of  tlu-  IiI.hmI  iVom  incr.-asinsr.     IViIiai. 

Ilic  icasoM    f..r  111.'   inaMlity  of  tlir  kidney  to  cxcn-le  mic  acid  midi! 

lii's  in  til.-  I'aet  tliat  its  salts  arc  amon-  tlic  l.'ast  soluMo  of  those  in  tl, 

'"''"'"■     '»  ■'^  ""  '1''^  '•" "lit  tliat  when  the  urine  ( Is,  a  red  sediment  ..; 

urates  eontainin-r  certain  i)ifinicnts  often  separates  out. 

The  nric  a.'i.j  „f  the  urine  is  possildy  derived  entirely  from  the  puri.r 

liH'tal)olism  of  the  Im,Iy,  i,,  \vhi.-li  tlie  nueleins  either  of  the  body  eells  „ 

of  the  ex.,-.'nous  f j  lak,.  ,,art.     It  is  decreased  durintf  starvation  an. I 

inereas,..!    l,y    caiinn:    f,„„I    rich    in    nueh.ins,    sneh    as    liver   and    swct 

lireads. 

rndcr  .ndinary  conditions  the  excretion  of  uric  acid  amounts  to  from 

<•:!  to   !•_'  yni.  per  day  ((1.02  to  (UO  per  cMit),  the  variation   l.ein-  dr 

'"•'"''•'"    "I ""'  ^t'l"'   "f  li-'i'llh,   .liet,   or  p.Msonal    idiosyncrasv.      Tl,. 

•''"'"1   "I"  a    normal    individmd   contains  mi   tlie  avera-e   1.8   mj,'.  "of  uri.' 

acid    per    11)0   e.c.      The    kidneys   are    ther..fore    al.le    to    concentrate   11,. 

'inc  aci.l  in  the  urine  from  :!()  to  (10  tim..s  over  its  concentration  in  tl,. 

hlood  i)lasnia. 

The  i.urines   found   in   colTee  and    tea    CcalTcine,  etc.^    are  excreted    in 

the  urine  as  salts  not  of  uri<.  acid  hut  of  methylated  xanthines. 
Hippuric  Acid.     This  is  a   constant   constituent   of  the   urine   of  her 

''''"'•"""   ■■"ii""i'^ 1    is   usually    present    in    small    annuints    in    human 

ii'-nie.     The  ann.unt   rarely  exceds  0.7  trm.  a  day,  hut  on  a  diet  rich  ii 

fruits  ami  veiretahles  it  may  ex 1  •_'  ..'m.     It   is  interesting    since  it  is 

th.'  only  urinary  nnistituent  that  is  synthesized  ),y  the  renal  eells. 
Amino  acida  are  always  i.resent   in  small  amounts  in  the  urine    eon 

"'"""'"-■  " "'I'"*-'  <"  •>•  !>•  Van  Slyhe.  dmnt   l.",  per  cent  of  the't.,tal 

nitro-en.     The  estimatimi   of  the  aniiiM.-acid   nitrofjen   of  the  urine  ha- 

not  li(.en  found  to  1 f  any  elinical   si.jnilieanee." 

The  aromatic  oxyacids  are  normally  present  in  the  urine  in  vnryiri' 
amounts.  TlM'se  in<.|ude  ph,.nn|,  indoxyl.  skatoxyl.  and  phcnyhiceti.. 
l)araoxy|.henyl.  proj. ionic,  oxynmndelic  and  homo-rcntisie  acids.  Tlies. 
ho.liesare  derive.]  fn.m  phenylamino  acids,  such  as  tvrosine.  trvptophanr 
and  phenylalanine.  It  is  l,eli,.ved  that  the  putrefactive  deconipositio, 
ol  in'otenis  in  the  larire  intestine  results  in  the  i)roduction  of  these  toxL 
hodies.  The  l.ody  prot.'cts  its.'lf  liy  oxidi/in;;  them  and  uuilin!,'  thet,, 
to  sulphuric  acid  to  form  the  ethereal  or  conjuu'aled  .sidphates,'^ whi,'! 
are  fotin.l  in  the  urine  iu  the  form  of  sodium  or  potas.sium  salt's.  Th. 
determination  of  the  amounts  of  these  hodi,.s  in  the  urine  has  therefoi. 
heen  taken  as  an  index  of  the  imtrefaclion  iroiuir  un  within  the  I.oue! 
The  chi..f  .if  th.'s..  I„»li,.s  is  urinary  iiiilii;i,i,  wh'u-h  is  f.iund  usually  ;i- 
a   i)ot!,.,.>ium  salt.     Th.'  t..st    lor  in.licaii   in  the  urine  e.uisists  in   oxidiz 


AMOr\T.    COMI'OSITION'.    ANH    CHAKACTKR    (IF    IKIM; 


r.:u 


ins;  tlio  iiidiixyl  in  nil  Mciil  suliifuiii  liv  means  of  iVri-ic  clilni-idc  to  indigo 
liluc.  iiiid  sliiikin<;  oiil  tin'  indiijo  liiuf  with  cldorofonn.  Tlio  depth  of 
!lie  eolor  of  tlie  ehloiofoi m  iilToids  a  rou^h  means  of  determining 
the  amount  of  indiean  present.  Tiie  faet  tlnit  the  indican  test  is  nega- 
live  must  not  be  taken  to  mean  that  tlie  intestinal  proeesses  arc  normal. 
lor  if  the  intestine  fails  to  contain  |(henyl,iied  amino  aeids,  or  the  proper 
I'Mcteria  are  not  i<resent.  no  indiean  will  lie  found.  On  the  other  liaml, 
I  lie  i)utrei'a<-ti\e  jiroeess  of  the  lary:e  liowcl  may  not  he  very  extensive, 
yet  the  annnnit  of  indiean  in  the  urine  he  increased,  heeause  of  greater 
.ilisorption  due  to  constii)ation. 

Skntoh,  a  fecal-smelling  sulistance,  is  formed  !>y  certain  kinds  of  bac- 
teria. The  grciiter  propoi'tion  of  this  substtince  is  excreted  by  the  liowcl, 
liiit  if  the  jieison  is  <'onstii)!ited,  sonu>  of  it  may  find  its  way  into  the 
lildod  to  imjiart  a  feeal  odor  to  the  breath  and  urine.  Its  prci^ienec 
therefore  has  s<inu'  diiignostic  importance. 

A  very  interesting  bocly  which  is  sometimes  fouml  in  the  urine  is 
I'lininnnitisic  arid.  It  is  thought  to  bo  an  intermeditite  step  in  the  metab- 
iiljsm  of  tyrosine,  and  is  found  in  the  urine  of  peoi)le  siifTei'ing  from 
nlkaptonuria.  The  disease  is  remarkable  in  that  it  ajijiears  to  run  in 
iMThilics  and  jirodiii-es  no  ill  ciTccts.  llomogentisie  acid  is  a  strong 
!i'<]ueing  agent.  an<l  for  this  leason  may  be  confused  with  sugar  in 
I'ehling's  test. 

Till    inon/inili-  rnn.itit units  of  the  urine  include  the  aeids:   chlorides, 
ulpliates  and  phosphates;  and  the  bases:  sodium.  potassi\nn,  magnesium. 
.iiid  calciiMii. 

The  Acids  of  the  Urine. — The  rhloriihs  compose  the  bulk  of  the  acid 
rndicles  in  the  uriin-.  Although  they  ajtix-ai'  to  be  necessary  constituents 
n(  tlie  living  cell,  they  do  not.  so  far  as  known,  enter  into  combinations 
'.\ith  the  organic  constituents.  The  tissues  a]ipear  to  re<iuii'e  a  rjither 
.leflnitc   concentration   of  sodium   chloride    in   order   to   carry   on   their 

iirk.  for  reduction  in  the  sodium-chloride  intake  of  the  body  results 
ill  a  reduction  in  the  chloride  excretion  by  the  urine.  In  salt  starvation 
the  chlorides  Jiiay  dis;ippear  entirely  from  the  urine,  the  amount  of 
'hloride  excreted  appearing  to  be  closely  relatecl  to  the  amount  of  stilt 
iiiirested.  \Yhen  the  inttike  is  constant,  the  rate  of  excieti<in  is  likewise 
iiKirc  or  less  constant,  but  !i  su<lden  reduction  in  the  stilt  of  the  diet  may 
i'c  accompanied  by  a  slight  decrease  in  the  salt  content  of  the  blood. 
^' ith  an  attendant  loss  of  water.  On  the  other  hand,  when  the  salt  is 
^'-'aiii  taken,  there  is  a  retention  of  .stilt  and  of  water,  with  a  coiiseciuent 
Mcrcase  in  body  weight,  until  equilibrium  is  re-established  on  the  old 
:  -.('1.  While  the  .ibove  is  the  Usual  reaction,  a  coii>iderii1)lf  letentiou  of 
-;iit  without  tin  increase  in  the  water  content  of  the  body  may  occur  in 


Ml' 


.    'f, 


i  r  ^ 


'I    ^  I 


ir 


1.  «     :! 


Ill 


!,!! 


!-*:f 


•'•'-  Till      I  \(  Id   I'MiN    HI'    IKINi: 

sDliip  npiJilli'lltlv     licinil.il    I'.-iMS.      This    is  iliii'    |)inli;ilil,\     to    llic   dipiisil  i,. 
Ill'  salt   ill  tlio  tissues, 

Carci'iil  stuilics  fail  t.i  (•(niliriii  ilic  idea  lliat   tlicrc  is  a  llxcd   iclatii.^ 
shi])  between  tiie  salt  ami  llie  water  of  tlie  Iwxly.      As  with   the  liitrou. 
lions  constitlieiits.   lio\\e\-er,   there   ai)l>ears   to    lie   a    relat  ions!ii|)   Ijclwei. 
the  rate  of  exeii'tioii  of  eliloiiiles  ami  tile  aimiiint  of  ehloriili'  in  the  lii.io.l 
Aiiiliarij  lpelie\es  that  this  relationship,  like  tiiat  of  the  exer.'tion  of  nir.i 
to    tjie    lilooil    nrea,    is   I'ajialile    of    lieinir   I'Spiesseil    mat  heinatieallv    i  si- 
l'a;;e   ■<-'•.    if  allowam-e    is   niaile    for   the    i';!,-!    that    Nat 'I    is   not    exeri'tci; 
after  it   falls  Ix-low   a  eertain  eoneeiitration   in  the  Mooil  e((iial  to  alioir 
'>.(i2  Kill,   per  10(1(1  e.e.     Tliis  level    is  more  or  less  ec.nslant    for  noini;i 
imli\  idlials,  hut   is  eonsideralily  iiiereased  in  diseasi'  of  the  kidney.     'I'lii- 
is  knoAMi   as  tlie   thresliold  of  eliloride  exeretion. 

The  aimiUlif  of  sodium  eliloride  exereted  in  the  urine  in  twi-nty-foni 
hours  varies  lietwcen  S  and  20  uiii.  a  da\ ,  aeecu-dini,'  to  the  intake.  T 
is  tliorefore  ai)i)arent  that  the  kidney  is  aide  to  eoneenlrate  tlie  sail- 
of  the  ))lasiiia  from  ten  to  twenty  times. 

The    Sulphates.     Sinee    the    inoriranie    sulphates    <lo    not     form    an    im 
portalit  i-onstitilelit  of  tlii'   food,  the  >rreater  |)<nlion   of  the  sulphates  n' 
''"'  II'      •       I'f  derived    from   tlie  sulphur   found   in   the   protein    mnlecnii 
l''or  tti,.-s      asoii  the  sul|ihates  of  the  urine,  like  the  nitrogen,  are  a  iiien 
lire   (d'   protein    metaliolism.      An    iiu'iease    in    the   nitroireii    excretion    i- 
aeecuiipaiiied   l.y    all    iiK-rease   in    the   sulphur   exeretion.    the    ratio   heinv 
about  .")  to  1.     Tne  daily  oii1|>iit  of  sulphur  is  lietween   I  and  :>  mil.     Tin 
greatest   output    is  in   the   form   of  the  alkaline  sul|iliates.  alioiit    10   p, 
eent   in  eondpination  with  aromatie  bodies,  and  a  small   amount   in  eom 
liination   wi;|i  amino  acids  and  neutral  iMi;anic  s.-ilts. 

The  iilKtsfihiitis  of  the  urine  are  derived  from  the  food  and  from  tl.' 
oxidation  of  phosphoriis-containiiifj;  bodies  in  the  tissues  such  ■.\- 
inieleiii,  lecithin,  etc.  The  daily  exeri^tion  \aries  between  1  and  ">  i;iii  , 
cab'iilateil  ;is  l'<».  When  calcium  or  mauiiesiniii  is  present  in  th' 
fooil.  the.\  are  excreted  by  the  bowel  -is  phosphate.  ;ind  proportionate! 
les>   is    found    in    the   urine.      Tl-  "ml    nsuall\    excreted    in    the    I'ece 

e(|nals  about  :'>o  per  cent  of  the 

Since  phosphates  in  the  urine  exist  as  a  mixture  of  the  mono-  and  iV 
sodium    hydrogen   idiosiihates,   they   have   an    iiiiiinrfanf    bearintr   on    li' 

reaction   of  the    urine,  the   ainonnt    of  ejo'li    varyiiiu'   witti    the   dci;ri 

the  acidity  of  the  urine. 

On  a  bea\,\  protein  diet  the  urine  is  acid  or.  .account  of  the  sulphiiri 
and  other  aci<ls  formed  frmii  the  im>al.  and  in  this  case  there  is  a  ^real' 
amount  ot  phosphoric  aeej  and  the  moiio-sodium  hydroiren  ])lios|>hat 
When  the  urine  is  alkaline  or  less  acid,  as  it   is  mi  a  vcjctable  diet,  tlier 


AMOINT.    COMI'OMTIOV.    AM)    (  IIAIIAI    IIU    (IK    1   HISi; 


y.v.] 


is  ii  liiifTc  iiiiiDiint  nl'  ilic  iliMMliiiiii  liyili(>(;cii  plmspliati'.  Sitii-c  ciileiiiiii 
iiul  iiiJiLriit'Niiiiii   iiliosiiliiitcs  iiif   iimrc  sdlnlilc   tlifiii   tin-   ilipliosjjliatcs  "f 

'111'  siinii'  inctiils.  ilc'piisits  nf  tl iirtliy   pliiispliiiti's  iiic  iil'tiMi  fuuiul  in 

iiiuliiil  nr  iill<;iliiH'  iiii.K's.  Wlicii  tlic  mini'  is  licjitcil,  tlic  dipliosplintc 
m|'  ciilcimii  Ini/ihs  up  intu  tiic  inniKi-cjilciiiin  iiml  ii  tri-culcium  jjlios 
filiMtc,  wliidi  aiM-diiiils  fur  ilii'  fiiii'  1iii'l)i(lit\'  ot'tcn  tiiki'ii  I'nr  alldiiiiiii  in 
tlir  ll.iiiic  ti'st.  Additiciii  111'  ai'iij  will  i-iiiisi-  tliis  to  ilisai)])!','!!'.  Tlic  crys- 
tiils  of  tiiplc  piiiispliatcs  uiiirii  oriMii'  in  jilkiiliiic  urine  arc  aMinioiiiimi 
iiiaLMicNiiiiii   plmspliati'    NII,Ml'I'()|. 


,   ( 


l:.,l,|:n-.l.    A.    P.:      i;...iil    .\.K:iim 

llMii;. 

I  i.-liiiv,   A.  }!.:      S,  11,1  Mill   111'   Tl  ill 


KiDNKV  i{i:ri;i{i:\(Hs 

I  .Miinoiriaplis  1 

I      l'!l\-iMl,,:;V,      I... 1,^1 w,     CliTll     &     Co..      I, Inn, 


mills,  (in  ,'M   \    I-,,,,   l...|if|i,ii,   liM7. 


lOriirinal    I'aprisi 


i:i|  I.  I\\\\ii,  H,  .■ii.",. 


Mi.Mlif.  T.  (i.,  nil. I   Miirkiiizir.  .1.  .1.:      I'n..-.   i;.,v . 

'•|i-liiiy,  A.  i;.:     S,.,ri'tiiin  of  I'riin'.   I!il7,  |i.   |s. 

\iMis';nii|  \V;iliiii.-il.i':     .loin-.  Ilinl.  ('In  ni..   liHi;,   \\i\.  L'n:;. 

M.  -piitli!il,  II.  ().:      Anil.   Int.   Mr.l..   l:il.-|.  :o  i.  ;:::;. 

\':I.:m-.I    nil. I    Will:      l'liv>i,.l..^l,.    ii,,iii,Mlr    .1     |.:ii  lH.I,i^i.|iir    ,lrs    niiis.    I'iiris,    IHM. 
.1,   P..   Iliiilliiiv  ct  til-'. 
•  \hi.!,.:,ii.   V.  C:      .h<:ir.    KsfLi.    \|.-.|..    I'.i|."i.   wii.  UlL'. 

rii:i~r  Mil.!   MrvriM     .1,    ir.  .\iii.   .Mr. I.   As-ii..   mil.,  Iwii,  :i:;i. 

\:iii  SlyUr,   D.'  I).,  nil. I    Mry.r.   C.    M.: r.    Hi.. I.  I'li.in.,    IIUJ.   \li,  r,W>:   am!    H'l". 

\v  i,  I'.iT.  L'l.'.  .■iihl  :;:'.l. 
■Knoulli.n.  r.  P.:     .Imir.  Pliv>i,.l..  I'.HI,  \tiii.  L'I'.i. 

I '.11 1-.. I.. ft.  .1..  iiii.l   Stiiiiil..   II.;      .I..1I1.    I'li.'.sii.l..    I'll.'.   \li,    I  l.-|. 
'I.'.nvnt  ii'c  .'inil   (loi'nalil  v:      .li.iir.   I'linnii.  tin. I    lA|..'r.  'riiir.ii..,   liMO,   i,  .-|7!l. 

A-^hiT  iiii.l  Pi.;iiTi-,  l;.  (■;.:     Z,.itv,-lir.  f.  Hi.. I..  l:ii::,  Kiii,  s:;. 

I'.'.'ii-.-r,   It.  ('•.,  nil. I  I'nrl.v,   i:.   I'.:      Am.  .Ii.iii.   riiy>i..l..   r.il'..  \v\\iii,  ::.'il1. 


[H 


:l 


JM 


,  I 


iv^    ■  < 


I'AKT  VII 
MKTAHOLISM 


U    -I 
u    '     t  . 


':i  •     I 


CIIAl'TKU  LX 
MKTAMOMSAr 

Introductory.     The  olgort  of  (liirostimi,  ;is  we  liavc  seen,  is  to  itihIii 
the  1'immI  cjiiiiililc  ot'  iili.sorption  into  the  ciiviilatory  lluiils     the  l)loo(l  nud 
Ivniph.     The  alisoibcd  I'niul  iinidiicts  arc  tlu'ii  traiispoitcil  to  tlic  varioii-. 
oi<raiis  aiul   tis.>'llcs  of  tlu'   l)o.ly,   wliciv   they   may   l>i'  cither  usee]   at   oiic 
or  stored  away  afxains,    future   re(|uireineiits.     After  beiu'r  used,   eertaiii 
substaiiees  are  i)rodueed  from  tile  foods  as  wa.ste  produets,  and  these  p:l^^ 
haelv  into  tlie  1)Io(m1  to  lie  earried  to  the  ortians  of  excretion,  i>y  which  tlie\ 
are  expelled   from  the  liody.     liy  compaiison  of  the  amount  of  these  c\ 
eretory  products  with  that  of  the  constituents  of  food,  we  can  tell   hnu 
much  of  the  latter  has  lieen  retained  in  the  Iiody,  or  lost   from  it.     Thi^ 
constitutes  the  suh.jeet  of  ;/<  ti< ml  witiiholism.    On  the  other  hand,  we  m.iy 
direct  our  attention,  not  to  the  lialancc  between  intake  and  output,  but  t  ■ 
the  ehcmic.il  chan-res  throujrh  which  each  of  the  foodstuffs  must  pa.ss  be 
tweeii  absorption  and  excretion.    This  is  the  subject  of  sprdnl  victahoHsui 
Tn  the  one  c.ise  we  content  ourselves  with  si  coniiiarison  of  the  raw  m.i 
terial  aeipiired  and  the  finished  j)roduct  produced  by  the  animal  factory 
in  the  other  we  seek  to  learn  somethini,'  of  the  particular  chanjrcs  to  whi.!; 
each  crude  product  is  subjected  before  it  can  be  used   for  the  purjuise  of 
driviuR  the  machinery  of  life  or  <.'  lepairinir  the  worn-out  iiarts  of  tlii 
l)ody. 

In  drawintr  up  a  balance  sheet  of  cener.il  metabolism,  we  must  select 
for  comparison  substances  that  are  eonunon  to  both  intake  and  output.  !■ 
freneral  the  intakn  comprises,  besides  oxyjren,  the  proteins,  fats  and  c,ir 
li()b.\<lrates;  and  the  output,  eaibon  dioxide,  water  and  the  vai'ious  TiitroL' 
enous  constituents  of  urine.  This  dissimilarity  in  chemical  structure  ]>,■ 
tween  the  substances  in^'csted  and  those  excreted  limits  us,  in  balancin'_'  tli' 
one  airainst  t'  nther.  to  a  comparison  of  the  smidlest  frafrments  into  wlii.  ! 
each  can  be  .ken  by  cliemical  asrencies.  These  are  the  elements,  and  o' 
them  carbon  and  nitro<rcii  are  the  ordy  ones  which  it  is  possible  to  measup 

,-.•!  t 


fl' 


Ml.TAIKiI.ISM 


535 


uidi  ai'curiicy  in  Itntli  iiitiikc  nml  (iut|iiit.  Kioin  Itiiliiiicc  sheets  (if  intake 
;iii<l  (iiitpiit  (if  ('ai'lidii  and  nitm^rcn  and  frimi  infuriiialinn  (il.taineii  liy  oli- 
Ncrvinsr  the  ratici  lietwcen  llie  aminints  nf  (ixv^'cii  cunsiniied  hy  the  animal 
.iliil  (if  earlii.nic  acid  excreted,  we  can  draw  fai-rcaciiini;  eiincliisidiis  re- 
LMi'dinfr  the  rehitive  ninmnits  (if  inntein,  fat  and  earlmhydi-afe  that  have 
been  iiivdlved  in  'he  inetalidliMii. 

As  has  already  iieeii  stated,  tiie  essenlial  natnre  of  ttic  nietalmlie  pnic- 
(ss  in  animals  is  ime  df  (ixidatinn  that  is.  nne  liy  wliieli  laiije  nnstalile 
iiMileeides  ai'e  iiniken  ddwn  tii  tlidse  tliat  ai'e  simple  and  stalile.  Diir- 
iiiC  this  pi'dcess  df  (iil(ihi)lis)ii,  as  it  is  called,  the  putential  en(>i-iry  lucked 
iiway  ill  the  larc;e  iiidleeiiles  liecdiiies  lilierated  as  actual  (ir  kinetic  enei'try — 
!hat  is,  as  iiidveinent  and  heat.  It  tliei-efdi-(>  liccdines  (if  impdi'tanee  to 
iciinparo  the  actual  eneryry  which  an  aiiiiiial  expends  in  a  irivcn  time  with 
I  he  eiieffr.v  which  has  ineaiiwhile  liecii  rendered  availalile  li.v  jiietaholisni. 
We  shall  first  (if  all  consider  this  so-called  <  \i<  rijii  Ixilintrr  and  then  pro- 
ceed to  ('xamiiic  sdinewhat  iiKire  in  detail  the  iinihrlnl  hiihnur  of  thr  Ixxhi. 

ENERGY  BALANCE 

The  unit  of  enor^.v  is  the  larpje  calorie  (written  ('.>,  which  is  the  amount 
nf  heat  rofpiircd  to  raise  the  temperature  of  one  kilofrram  of  water  throu^rh 


tio  (U>}rrco  (('entifirade')   of  tem|)erati 


\V( 


•an  determine  the  calorie 


\,iIuo  by  allowing;  a  measured  (piaiitity  of  a  suiistance  to  liiirii  in  com- 
presypd  oxysrcn  in  a  steel  liomh  (ilaced  in  a  known  volume  of  water  at  a 
'irtain  toiiiperature.  Whenever  cdinliustidii  is  coniiileted.  we  find  nut 
IliroUdh  how  many  de<rrees  the  temperature  df  the  water  has  li(V(ime 
iiiis(Ml  and  multiply  this  liy  the  volume  of  water  in  liters.  Measnrt^d 
ill  such  a  ciiloriHK  tir.  as  this  apparatus  is  ealleil,  it  has  lieeii  found  that 
the  liuniher  of  calories  lilierated  liy  liurninir  one  trrani  of  each  of  the  proxi- 
nate  principles  of  fiKid  is  as  follows: 


;o       ii 


!i'  icr 


it 


'    ', 


I 


m 


M 


rilll."livcli;iti 


(Sliinli    t.l 


■t.o 


T'inti'in 
Kilt    .  . . 


Th 


e  same  nuiiiiier 


her  of  calories  will  lie  lilieratecl  at  whatever  rate  the  coin- 


istidii    priiceeds.   priivided    it    results   in   the   same  end    pi'ddilets.      When 
ncli  as  supir  (ir  fat,  is  liurned  in  the  presence  df  dXVLren,  it 


suiistance,  s 


rid 


s  ea 


rliiin   dioxide  and  water,  which  are  alsd  the  end   iirddiiets  (if  the 
■m  of  these  f IstnlTs  in  the  animal  liody  ;  therefore,  when  a  srram 


I'taholi 


!'  suirar  or  fat   is  (lui' 


•kh'  liurned 


;i   calorimeter,    it    releas(>s   the 


'The  .li^liiuti 
.it  ciut^y  is  IieifiR  Ri" 


.1. 


'ipi  latnrt'   itm^l    tir   i  If 


Ittv  of  ,litual    llfil'   (■' 


K>  ;  tl  f  lattt T   nil  rtiv  I« 


iIk 


■r.l I       Th. 

at   wliuh  tin 


I. 


•,    i'l 


!  ] 


i 


■l^.,;i 


■l\ 


% 


:.:Hi 


Ml  TM'.OI.ISM 


•liiioillit  111'  iiii  l'i;,\  .!>  \\llcli  it  is  «^1(H\I\  ii\iili/('il  ill  the  ;||ii|ii;i|  lii.cly.  Hui 
llir  ci-c  is  ilirrcii'iii  r,,!-  inulfins,  hci-jUlM'  tlii'si'  yield  less  ciiiiililctcly  hm 
'li-'i'il  "11,1  piHihiri,  ill  till'  iiiiiin.'il  IiimIv  tliMii  llicy  \  ii-M  when  Imnicj  ii 
nvvL'i'n  :   --(I   tliiil.    Ill   ii^i'i'l'tilili    ll;e    |ili.\  siiildiric   eiiefy^y    Viiluo   ot'   liriitcili,    ^\ 

liil"<'    'iedlh-I    n ils    |i||\siriil    li.'iit    \;iliie   tile   |ili\sic;il    liciit    VmIiIc   nt'   tl 

ill! |>l(tr|\    ii\i,li/|.il   eii<|    piiiclllets  «<i   its   iiiel.-iliulislii.      it    is  (ilivimis   ||i;/ 

\\i'  ran  riiiii|iiile  ill.'  tiital  iivailiil'li'  ciiiT^ry  (if  (lui-  diet  liy  iiiiilli|ilyiiit.'  tl  ■ 
'|Uaiitil.\   nf  earli   !'(  h  ii  l-I  iill'  liy   it-  caldi-ic  value. 

Methods.      In   duler   l.,   mc'isme   t' iicmy   that    is  aetiiallv    lilieiat.-.i 


l-K     l.t        K.v|,,,., ,    ,,,|,.,,,n>l,r    ..1    ,l„.  l<.,,-,l:    >,,„,.    Inviituu     ..1    r.,t|...|,,„v.    IScUcnu     II, -|i; 

N.iv     'l-^^^      .\t    II,     ■itlil    I-    -,,n    ,1h     k,1,|,  «,,h    the    ,,l.s..,  [.Ihin    ^,ll.,^;    .nil, I    in    |hc    miil.ll,-.    at    1 

Unk.  I!u-  ,-,,,lru-  ,..i;,,,,l    |.,M,     l.r  l ,  i;-:!,-iliiii;  ih,-    I,  Hl|i,  rallir,-  ,.f    t!i,-  ,l,,iiMr    «;il)<  of  thr   in|,.nm,t, 

■\'    '!■     ,^'l,m,    1.  ••    ,.    il,.    ,.    M^,,,    ,vI,,hI,..  , I. :,-;,■-   S. ;,-,..,■    ,,    \  ,(,,(,    I!  . 

'"   ''"'   ■■" '■'!    '""1.^.   "I     liiil-l    alsn   lis,'  .1   caliH-iliielei-,   hut  ,.l"  soliiewhat    ilil 

1',  r,'iit  r,,iisliii,-tiiiii  iVinii  Ih.-it  iise,l  h,\    the  eheiiiist.  I'.ii'  we  have  tii  itn.vi.l. 
r,,|'  hill-  e.iiitiinied  nhservatii'iis  and    I'.'f  ,iii    luiiiiternipted  siii>|i|y  nl"  ux' 
-' i>   1"  'lit-  iuiiin.il.      Aiuiiinl  (,ilo,:,in  h  r<  are  alsd   usually    pnivided    uil' 
means  I'ur  the  iiieasureiiient  nt'  the  aiiinuiits  i.t'  earlMiu  dinNide  (and  watr, 
■  lisehai-ued  and  nf  dxytren  ah.sdrhed  liy  the  .iniiiial  during,'  the  i>l.sei'v;iti,,i 

Siieh   ivs|,i;'ati,,n  ealnriiiietei  s  ha^'  I n  inade  f,.>'  a!!  sni'ls  i.f  aniiHaN,  >' 

iiiiist  peifei't  \\n-  use  (in  man  liaviii}.'  heen  eoiistrueted  in  Anieriea   (see  Fij 
!7t>.      As  iljusti'atilii:  the  e.xtteine  aeeur,ie_\   111'  even  the  laiji'est   iif  tiles, 


Ml  rM;n|.|>\| 


337 


i.s  iiiti'i'i-Ktiiiii   ti'   iM>t('   tluii    till'  ^ii'tii.'il    liiMt    ''JMii   iiiit    wlirii   a   ili'llnitt' 


iriii   cxiicllv   iMiiTt'StHiii 


.iMiiiiiiit  lit'  alciilnil   III'  itlici'  is  liiii'iii'il  ill  niic  nt'  tl 

III  the  iimoiiiit   iis   iHiasiiri'il   liv  tilt-  siMiillcr  IhmuIi  i-aloiiini'tcr.      All   (if  tlic 

.  llc'l'(.'V  liliiTalc'il    III   the  ImmU    i|()i'>   licit.   hnWiMT.  take   tilt'    liillll   (if  liciit.      A 


lillilc  iiliiiilliil  a|i|i('ai's  as  iii('( 


lianical  wniU.  sn  tiial  In  iiiimmihi  in  cali 


ill  (if  the  ciiciuN    that  ai 


II  annual  cnih'iiiIs.  i  ih'  m 


list  ;iili|   In  till'  actual  cal- 


iirics  j.'ivt'ii   nut.   llic  calnric  i'i|iiivali'nt   ni'  tlic-   miiNi'iilai'   unrU   wliicli   lias 
lii'cii  pcffdniii'il  liy  the  animal  diirinir  the  iicfinil  nf  nlisi'i'vatinn.     'I'lii.s  can 


nicasiii'cil    li\    iiicaiis  nf   ai 


1   ci'i.'niiictcr.   a   calnii 


(■   (•  ii'i'i'-piin 


lill'J'    tn    4.'J' 

kildSi'iini*  iiicli  IS  (if  udi'k.  That  it  lias  ln'cii  |inssili|(>  tn  strike  an  acciirati 
iialaiicc  lictwccii  the  intake  aliil  tlic  nut|iut  nf  eiiei'i.'y  nf  tlie  animal  lindv 
is   nllc  (if   tlic   aclii('\eliielits   nf   iiinilein    cNpeliliieiital    lii(iln(.'V.      It    call    Ik 


iiic  ill  the  case  u\'  the  human 


animal  :  thus,  a  man  dniiitr  wnrk  nii  a  liicvc 


.'I'uniiictel'   in   the   Bcneilict    calnrimetci'   umnc  niit   as  actual    heat   4.s:!.">  ( '.. 
iiiiil  (!i(l  wink  e(|ualliiii_'  li"-  ' '..  '.'iviiiir  a  Intal  nf  ."i.t:!,")  ( '.     Ity  draw  inn  "I' 

,1   lialaiiee   slicel    nf   his   intake   and    niit|iiit    nf    f I    material    durin<r   this 

l>iriiid,   it   was  fniiiid   that   tlii>  man    had   cnnvuincil   an   amniint   eapahlo  nf 

weldiiijr  ."i.4."i!l  ( '..  whieli  ma,\   he  ( sidcrcd  as  exactly  lialanciny;  the  actual 

nutput. 

It    wnllld    lie   nut    i>(    place    In   'ji\e    a    full    descliptinn    nf   the    I'cspirat 


ilnrilllcter    here.      'I'lie   ircnel'i 


inn 

il-IIUcImn    will    be   seen    Irnill    the   acenm- 


l';ili.\  ill'^'  li^urc  nf  the  fnrm  nf  ,'ipparatus  in  use  \'<u-  palicllts  in  the  Russell 
Saiio  Institute.  New  ^'ork.  <  »nc  nf  the  must  intereslini.'  details  nf  its  cnii- 
•  'niitinn   cnliecl'lis  the  means  taki'ii   tn  prevent   any   Inss  nf  heat    fi'niii   the 

,i|i'iimelei'  In  the  siirrnuiidiiiij'  .lir.     This  is  aeeiimplishcil  in  the  fnllnwin;; 

\:iyt  Tie  iiinei'mnst  layer  nf  the  wall  is  nf  enpper:  then,  sepafateil  IVnm 
'his  hy  an  air  space,   is  annthc'  wall  nf  cupper,  niitsidc  nf  uliieli  arc  twn 

V  linden  walls  separated  frnm  each  ntluT  ami  frnm  the  niiiiT  i-npper  walls 

■y  air  sjiaces.     The  tw pper  ualN  .in linected  thmiiLrli  thermneleel  rie 

■I'llfilcs.  sn  that  an  electric  eiirrcnt  is  sel  up  vvlieliever  tlier(>  is  an\-  difl'er- 
'  iicc  in   their   li'iiiperatures.      The  ciirreiil    is  nliserxed   liy    means  nf  .'i    ijal- 

'' .■inniiK^tcr  placed  niitsidc  the  calnrimeti'r.  and   finm   its  iiinM'inrnts  t! li- 

^|'l'\'cr  cither  heals    llji    nr   cnnls   dnw  n    the   nllter  Clipper   walls   sn  as   tn   iHiv- 

ci't  the  difference  nf  temperature  causiii!.'  ihc  current.     This  is  dnnt>  li.\-  an 

ilectric  heatinu'  device  nr  hy   cnld   water  tuhes  placed   lietwcen   the  niiter- 

nst   Clipper  and    the   innermnst    unndcn    walls.      Since  the  tempei'atili'e  nf 

■i,e  tun  Clipper  walls  i^  the  same.  Ihere  eaii  he  nn  e\clian'_'e  nf  heat   hctweeii 

'  ''111.  and  cniisi'ipienl  ly  nniie  I'f  the  heat  that   is  alisnilicil  Ii\    the  inner  enp- 

'1'  Walls   is  allnwed    In  lie  carried   awa>.      .\ll   the  heat    irixcn   niit   hy  the 

lillial    is    alisnlhed    ll\     llm    sile.llll    nf   cnid     Walcr    llnwihl;    lll|nl|oll    ijli'    enils 

A   l.il..i:i;iin    mil.,    i,   ih,-   i.i,..I;uc    .,i   Hi,    l.M.I    III    l,il,.^i,,„i,   ...-i!tii.li,  ,|    In    ilir   ,|i..,.,.     in   TMl.r^ 
'cll   wlil.li    It    i~   liftril 


H 


1  -f! 


ili^l- 


' '— "  •  * 


!; 


,    '  I 


i    '"I 


■  ' 


)  ^ 


I  I 


r.:iH 


MKTAIUIMSM 


of  |>i|H'  ill  tin-  cliiiiiilMr.     Tin-  licit   used  to  va|)iiii/.i'  the  iiidiHturi'  ti<.  i 
skill  iiinl  ImiiiH  iiiiisi  of  coiirsf  also  lie  iiiciisurcil.    'I'liis  is  i|oim>  liy  collii  : 
iiii;  the  wtitcr  viipor  in   a   Milpliiirir  ai'id   liottlc  [ilaci'd   in   tin-   vcntiln' 
ill},'  ninviit.      It.v   iiiiilti|ily iiifr  tiic  >.'iiiiiiN  of  wntiT  hv   the  factor  for  tl  . 
latent  heat  of  \  a|iori/atioii.   wi-  olitain  the  ralorics  of  lioat  so  ('liiniiiati  <i 

'The  caloriiiirtci ntaiiis  a  coiiifortalilf  linl  ami  is  (iroviilcd  with  !».. 

uindiiws,  a  sticif,  a  tclc|ilioiu..  a  fan,  a  TiKlit.  and  a  Howies  stetlioseo|)e  |..i 
eoiiiitini;  the  jiillse.     Tlie  onlinaiy  ex|>eriiiieiit  takes  aliont  as  loiif:  as  a  trip 
from  .\e»    York  to  New    London.     I'atieiits,  as  a   rule,  ilo/e  from  time  t.. 
time  or  else  I IV  to  work  out  some  seheme  liy  whieh  they  eaii  amuse  them 
selves  without  mie.iny.     After  three  or  four  hours  they  are  rather  lion.l 
liy    the  (|uiet,  and   the  oliservatioiis  are  not    pri)lon>t<'<l   iM'yond   this  tiiin 
They  are  allowed  to  turn  over  in  lied  oiiee  or  Iwiee  an  hour,  liut  readin.' 
and    tele|ilioiiiiit5    are    diseoiirairi-d,    since    these    iiierea.se    the    liietalioliMn. 
The  air  in  the  liox  is  fresh  and  pure,  the  patient  suffers  no  discomfort,  aiil 
oli.jeetioiis   to  the   procedure  are   very    infreipient.     .Most   of  the   paticM!^ 
are  only  too  <;lad  of  the  extra  attention,  and  they  insist  that  the  caloi 
iineti'r  has  a  niark<'d  theiaiieutic  value."     (  Dii  Hois.) 

Normal  Valuei.  -  Ifaviii}:   thus   s,itis(ied   ourselves   as   to   the   extreni' 

accuracy   of  the  method  for  ineasuriiit'  eiieriry  output,  we  shall   now  i 

siller  some  of  the  conditions  that  control  it.  To  .study  these  we  must  first 
of  all  deteriiiiiie  the  hasul  hmi  i,ri>ihitiii)ii — that  is,  the  smallest  ener^j;, 
ouf|<ut  that  is  compatilile  with  health.  This  is  ascertained  liy  allowiii};  .-. 
man  to  sleep  in  the  calorimeter  and  then  ineasurin<r  his  calorie  output 
while  he  is  still  restiii<;  in  lied  ill  the  niornin'r,  fifteen  hours  after  the  hi-' 
meal.  When  th,.  results  thus  ohtained  on  a  numlier  of  individuals  aiv 
calculated  so  m  to  repr'esent  the  calorie  output  per  kilofrrain  of  body  weijriii 
in  each  case,  it  will  lie  found  that    1   < '.  jicr  kilo  iier  hour  is  discharc  i 

that  is  to  .say,  the  total  enerj,'y  expenditure  in  24  hours  in  a  man  of  T'l 
kilos,  which  is  a  srood  avera^'e  weij:lit.  will  lie  70  X '-4    -  l.fiHO  ( '. 

When    food   is   taken   the   heat    priNliii'tion   i-i.ses,   the   increa.se   over  (!• 
Iia.sal   heat    production  aiiiouiitiiij;   for  an   ordinary  diet   to  almut    10  p. 
cent.     P.esides  lieiiii;  the  ulfim.itc  .source  of  all  the  lio<ly  heat,  food  is  thci. 
fore  a  direct  stimulant  of  heat   production.     This  sprrifir  ihpKitiiif  actim 
as  it  is  callcj,  is  not.  however,  the  same  for  all  groups  of  fisidstuffs,  licin 
■.'leatest    for  lu-otcins  and    least   for  carliohydriites.     Thus,   if  a   staiviii 
animal  kept  at  :!•".     ( '.  is  "^'ivcii  protein  with  a  calorie  vjiliie  which  is  e(|u 
to  the  calorie  output  durint:  starvation,  the  ejilorie  output  will  increase  1 
;iO  ]ier  cent,  whereas  with  carliohydrates  it   will   increase  liy  only  (i  1>' 
c-ent.      Kvidcntly.    then,    pr.itein    liherali's   inm-li    free   heat    diirinj;   its   ,i- 
simil.'tioii  in  the  animal  liody;  it  liurns  with  a  hotter  flame  than  fats  . 
carliohydrates,  althoufjh  liefore  it  is  completely  burned  it  may  not  yici  ' 


MITAIMII.IKM 


5:1!) 


N.  iiiiirli  cmrKy  "h  i^  the  (■ii.s4>,  I'm-  cxiiiniilc,  wlini  I'iitx  iiic  Imnicd.  Tliin 
I'ic'iiliiir  |iiii|i('it\  III'  iHiiti'liiN  iii'riiiiiits  t'lir  tlii'lr  wcll-kiinwii  liciiliii);  i|Uc'il- 
iiii's.  It  rxpliiiiis  why  |initi'iii  i'iiiii|iiim's  mi  \nr)ii'  a  |ii'ii|iiirtiiiii  of  the  ilit't  "f 
I  rn|ih's  liviiiif  ill  I'lihl  i(  t'ii  ii".  iiikI  \\\\\  it  in  cut  ihiwii  in  llic  ilict  of  thoHc 
uhotlui'll  iiritr  the  ti'o|>if.s.  I iji|i\  iiliiais  iiiaiiitiiiiicil  nil  a  low  pniti'in  diet 
iiiaN   NUfl'i'i'  iiitt'MM'l.N    friiiii  colli. 

If  wc  mill  to  the  liasal  heat  |iriiiliictioii  nf  l.tlSd  ( '.  another  KiS  ( '.  (nr 
III  per  cent)  mi  aeei.iiiit  of  fonil,  the  totiil  I.s4s  ( '.  neveithelesH  tails  far 
>liort  of  that  which  we  know  iiiiist  lie  lilieiateil  when  we  calciihite  the 
ivailalile  eliei).'.\  of  the  iliet,  which  we  may  take  as  ■_'.."((H»  ( '.  What  lie- 
■  oMies  of  the  extra  fuel  .'  The  answer  is  that  it  is  iiseil  t'or  miisi  iihir  iiorl;. 
Thus  it  has  liecn  fniiiiil  that  if  the  oliserveij  peison,  insteail  of  lyini;  ilowii 
111  the  calor-iiiieter.  is  iiiaile  to  sit  in  a  chair,  the  heat  proiliiction  is  rais^'il 
li\  H  per  cent,  or  if  he  pert'ornis  such  iiiovciiients  as  woiihl  lie  necessary  for 
"iiiinary  work  iwriiini;  at  a  ilesk)  it  may  rise  •_'!>  per  cent-  that  is  to  siiy, 
lo  !'(!  ( '.  per  hour.  There  is,  however,  practically  im  ilifl'crcnce  in  the  eii- 
'  r^ry  output  of  a  person  lyiii^'  Hat  or  lyint;  in  a  seini-reclininj^  posi- 
liiiii,  as  in  a  steamer  chair.  AlJowiiii.'  citrht  hours  for  sleep  ami  sixteen 
I'liiirs  for  work,  we  I'aii  ai-coiint  for  almi.t  ".MtiS  ( '.,  the  remainin;:  :i<MI  oijil 
I  .  that  arc  rei|uirei|  to  lirintr  the  total  to  that  which  we  know,  from  statis- 
tical tallies  of  the  iliets  of  such  woi-kers,  to  he  the  actual  ilaily  expenilitliro, 
i'ciinr  line  to  the  exercise  of  walkin;;.  If  the  exercise  is  more  strenuous, 
^lill  more  calories  will  lie  expemleil  :  thus,  to  asceml  a  hill  of  l.d.'iO  feet  at 
:lic  rate  of  L'.T  miles  an  hour  reipiiics  4(1"  extra  calories.  Field  workers 
niiiy  expend,  in  ■J4  hours,  almost  twice  as  many  calories  as  those  en^rajred 
i  1  sedentary  occupations. 

Standard  for  Comparison 

Will  II  the  eneriry  output  per  kilo  hnihi  in  niht  is  determined  in  animals 
I  \ar.\in<;  si/e.  the  values  are  irieater  the  liirhler  the  animal.  Tlii.s  is 
\i(lent   from  the  followin;;  results  olitained  on  do'rs: 


Hi 


I 


.  t     l!»* 


r   ]i 


1 1 )  :;i.j 

(J)  I  S.J 

(.■;i  iMi 

(  t  I  o.."> 

(,-)i  ."..I'.i 


Ilfiit   /iritiliirtitm    :/■    rtih>ii,s 
jM  r   l.ihi   jii  r   (III II 
.".."i.lis 
li5.L' 
tl.-|.ni 

(>i;.ii7 

.ss.ii* 

I  Iflll'lll'l  I 


When,  on  the  other  hand,  instead  of  liody  weijrlit,  the  area  of  the  siir- 
'cui  tiic  lioitv  is  lakeii  as  llic  tiasis  oi'  caii-iilal ion,  results  that  are  almost 
'iistant  are  olitained.  Followiii';  are  the  results  in  the  aliove  ani'nals  on 
lis  basis: 


L 


r 

u 

-:|':. 

.'   ; 

■ 

\ 

n    ' 


vio 


Ml  TMIiil.IsM 


1  I  1  lii.T.'iii 

'  J )  r.i'f.j 

(It  .•■.,:.' I 

.:.  I  -.Mj:; 


II'  ,il    /•nnlil,  hnn   r;<    i''hii  I' 

f',  ,  y,f>,,i,i   nil  ft  r  "C  *'<r- 

itiri'  fit  r  iht'i 

10.".ii 

m>' 

118.-1 

1ir.:! 

I  •-•  I  ■-■ 

•  irii.i,.., 


>cil-||    |.'M||l>    l|,|\r    |>lo||l|>t<  <l    illMt'MIV    |..    IMi||i-|l|l|c    lll.ll     llic    ilctrniiiiiii,' 
l''l''l"l'    ill    til.      imIiHIi'    nll!|i!|t     ,,r    u;|||||    l.lnnill.l    i|t|irilill>    is    till'    nhllin      m(, 

'''''    I'l"  till'  iiiiiiiijl.     Tills  Is  '/iTiilrr  till'  siiialliT  till'  iiiiiiiiiil.  with  tlic  i-"i 
Mi|iiciin'  tli;ii    111  ill    is  iiiiiiT   r;i|.i.ll\    Inst   \n  till'  Mirnimiilint;  nir   rium   |l,. 
siirriicf,  lliiis  !•  i|iiiiiii::  iiiuir  ,ii|i\c  nmiliiiNlioi..     I'litil  <|iiito  rcccnlly  it  li,,- 
Iwrii  ycliti;ill\    III  jii'V  I'll  ili.it  ..iicli  ii   ri'hltiiilisliii    liftwcni  Itiwly  siii'l'iirc  mipI 
liiiil   iHiiiiiiilinii  iliil  .iitii.illv  I'sist.  I. lit,  tlialiKs  III  till'  wiilk  of  !•'.  <!.   Itni. 
ilii'T  .iiiil  !■;,  !•'.  anil  l>    l>ii  |!iiis'\  it  is  nnw  kiinwii  lliiit  tin'  iMli'iilaliii,;>  \\i'\' 


f  I  1 


I' 


r!! 


!1.-  i 


I    s 


200 


180 

5160 
£150  -N 

(1,130 
120 


ICO 


20  30  40  -JO  60  70  80  00  100         110 


1 

i; 

■\ 

^ 

\ 

^l[ 

X, 

^., 

^^ 

g 

14 

„ 
\ 

'., 

\. 

\  ^ 

II 

N, 

\ 

\ 

\ 

\ 

\ 

X 

X 

\, 

^-^^ 

^ 

\\ 

\ 

> 

'°\ 

\, 

\ 

\ 

\ 

\ 

"- 

\, 

^ 

\ 

" 

"^q 

ir> 

^    \ 

N 

V 

\ 

\ 

N, 

X 

K 

V 

^ 

X 

> 

So 

\ 

\ 

\ 

s 

\ 

\, 

^ 

V 

\ 

\ 

\ 

'\ 

v 

1 

4 

Ii 

•fi' 

4- A. 

^ 

N  ^ 

'x 

"' 

\  ^ 

L 

\ 

kJO              II 

'00 
190 
i80 
170 
160 
150 
140 
130 
120 
lllO 
100 


I'O  30  40  50  60         70  80  90  100         HO 

WEIGHT-KILOGRAMS 

•■'*■       I""         il.nl     l-I     ■liUIlMi;n^     •   il  I.m  -      .i    .  ,,        !     lu.ii;     ;.     -ii.i!>'     nuKIs     l|..nt     UUwIil     111     - 
b:.ii    -    iWli    .ir-i    l.ik'lil    HI    .,  ni,,,i,  1,  I  ,    ilh    I    .1 In  i:    ]..    ih,     1..1  ikmI.i  :      Ana    lS.|.    Cin-I         W 

111';  Ml!    !•:•'  \ri.M     ii'i,.!!.  |i,,K.,i,  :n'.i  !''ii...i..   \t,h    I'll    M..!.,  i"ir,  V..1    ir  ■ 

liasiil    U|iiiii    iiii-.,i  II  i-t    i-ip|n|iillalin|is  ni'  llir   limly   siirrai'i".      In   tin-  oliirr   i< 

-' ■"■'•li''^  'I aii-iilalh.|i   \\.is  iiiailn  li\    ii^iii'^a    riiniillla   kllnUli  as  .Mi'i'll's.   : 

\'liinli  wci'.'lit   was  iiiiilli|ilii'i|  liy  a  ni-rlaiii   rai-lnr  i  \  i/..,   I2nil2  ■•  \'  Wt'lsjlit 

l»ii    lliii.s.   |i,p\M'\i  r.   lias  sliuMi   llial    an   a\rrai;('  itiui-  iil'    lli   |iim iil    is   ii 

niiriiil    ill    iisih;.'  !lji>,   I'm  nulla.      i''iii'  ; iiratr  iiiraMircini'iil    tin'   Imily    w.' 

luMiiil    uitli    thin    iliiilrruiai'.    wliii-li    was    tlnii    iiii|ircv'iiiit('il    uitii    iiirlli 
I-arattiii  ain!    ii-iiif'Ufi-tl   \i;!!i   |'a|ri-:-  strips  III  piTM-nf    it    i'niiii  chaiisrintr  i' 

ari'ii   wlirti   ii'iiinviil.     Tli;     i Ii'l   ul'  tlic  siirt'ari'   was  iiflcrwarfls  cut   ii 

ililii  (lat  pici-cs  ami   |ili.itiini-a|ilii  d  mi  |ia|ii'i'  nf  iiiiirunii  tliickiit'ss,  (lie  pa 


Ml  I  yi'.iil  l-M 


h'loMi    till'   rt'Mills   it 


ill 


■.V     111 


lilts  lii'iin;  tlii'ii  iiil   iiiil,  iiiiil   wii'.'lii 
.ili'iiliiti'  till'  iii'liMl  «iii  Nii'i'  ai'i'M. 

Wlll'li-    till'    hcMU'llt    illlll    \M'I);I.I    Jlr    I 
'I'   till'   MirCi Mil    III'    si'l'llllll    In     UMIllf    III,.    riillinMlii;    t'lllMlllhlH:        A         W'"' 

H"  •  71.>t:    .1    lii'ihi.'    till-    Mllli iiUM    III    M|i(iili'    ri'i|tillli'liM;<  ,    //    flif 


r.iiiK 


'iii'iiii'  i-oiii|Mituliiin 


I'iu'llt     III    c'i'MlilMltil  s  ,    illlll      II.     Illi 


«c'ii:lil    III    kiliiaiimis.      I'liisi'il   III!    this 


iil'lllllhl.    il    I'll, lit    |i;|v    III 

'  rniiiii'il  ill  ii  L'liii 


I'll    I'll!  Ii  <l    hill 

I'lLT,  n 


II    \t  Inrli    llir  olll'JM 


!■{•  .iliil    lilil\    lie  ilr- 


■'        Aiiiiilnr  hiiiliiiil  r iiilv    i'iM|i|iiM'il  li\ 

r.iMl'llil't    is   lljisiil    nil    IImmsIIM'MK'IiIn    lllillli'    Iriilll    |llilllllV  .illllis   llf    till-    >llli,ji'i'l 


III      \ai'llll|S     llllNIN. 


I(.\    till'    IIM'   n|'  thtsi'    iiini'i'  MiTilfiitc   Mii'ilMll'i'MirliN  iil'   ImhIv    siii  fai'i',    i| 


kiKPVMi  llijil.  jililiiiiiL'li   Ilii-  siirr;iiT  :iii  ;i  la 


U    V'l\is    IIS    rcili-t.ilil     liMllls 


iiir  I  III'  I'lirrL!^   Ill 


lljiiil  111'  ililTninl   iiiiliv  JiIim 


>l  ^iiiiiliii'  Illlll 


ilTrl" 


1>  il   inili'li   liioli'  iii'i'lHilli'  liilsis   liir  rii|ll|iillilil.'  Illiisi'  of  liilVi'trlll    laliiHiitiin 
iliilllills,   Ml    il    lilTiiUs  ijiiWIl    wlicli   il|i|.rir.l   ti II    ill    wiijcly   ilHrililiL'   stillis 

if  lidily  iiiilriti'iii.     'riiiis.  ill  t 
il 


II'  I'.isi'  III    a   iiiii 


II   « liii  stai'vi'il   (m-  A  ihiimHi. 
I'  calmir  iiiil|iul   |iri'  M(iian    mtti'i'  .if  si'il'ai'c  ili'iTrasi'il  touaiiU  tiu'  cinl 


■r  till'  fasl  li\   -Js 


M'l'  iTiit.      ( »|i\  Minv|\  . 
ill    I'l 


roll',  il   « Illlll]  lie  il ri'i't   II 


law  I'lilii'lllMuiis  ii;;ari|iini-  |i..>mIi!i   I'lialii:       III  i'iii'li:\   iillt|iiit  nf  a  M'l'irs  nf 
iiariati'il  or  rnriniiriil    iiiilu  iihials  li\    riiiii|iaiiMiii  nf  tlu-ir  ralnrii'  iiiil|iiit 
I'  si|ilar tcr  nf  siirfaii'  with  thai  nf  imniial   iiii|i\  iiliials. 


The   lilt. 


iiiiiMiiiL'    I'arliir   nf   i'iii'ry\    niitiiiit    is    iiinliiiilili'ilh    iln'   I'l'iicfal 


'iiililiiiii  llf  Imilily   iiiiti'ilinii     ilii'  ai'Tn 


r  mass  llf  i)riitii|ilasin   nf  tin'  limlv 


I'.i'licilii't  I.      Thai    llii'li'    is   a    lrlaliii||slii|i    Iirlwciii    llii'    Imily    siii  fi ami 

I'tallilllslll      is     llllllnlllltl  .1.      Illlt      till'      li'lalin|l>||i|i      is      lint      a      I'allsal      nlli'.         Al 

;ir«iMlt.     llli'i'ifnli'      till'    niilx     safr    liiillinil     In    i'iii|ilii\     in     ri ilii |ia fi IMJ;    till- 
'   ilalii  li^iii   i<\'  iinniial   ami  ilisrasi'il   imli\  iililals  is   that   ralliil   liy    ISi'lli'ilii't 

till'     li|'li|||l     llU'tllnil."     ill     whil'h     till'     Illi'talinlislll     li\     i_'liil||is    nf    IH'I'SIIIIS    nf 

!  1'  hiiu'Iit  ami  wri'^'ht  is  i'iiiii|iavi'.l,  it  InihL'  assiininl  thai  siH'li  iiiili\  iiliials 

•  i\i'  till'  satiii'  tri'iii  lal  ..'I'liuih  iilatinns.     I'"ni'  fli,.  aiiiiliiMtimi  <<(  tliis  (.fnip 

'lliiiil.  Imui'Mi'.  iiinii'  i'\tiii'i\i'  ilata  will  Im  riijuircil  than  I'xist  at  pri's- 

i.l.  aiiil  althnii'.'h  snim'  ni'  tin'  I'niii'liisiniis  ilrawii  frniii  fi'sults  I'ninpiiti'il  mi 

'1'  siii'f; aii'a  liasis  may  liavr  In  lir   ii'\  isnl.   it    is   lunlia'ili'   tliat    tlmy 

'•  ■    ill    'Ji'lli'l'al    rnlTi'i't, 

Influence  of  Age  and  Sex 

Till'  liii  i-^y  iiiitpiit  is  Inw    ill  till'  iii'ulx    lini'ii  ;  it    i  iii'ii'asi's  rapiilly  iliiriii^i- 

!r  lii'st   \i'ar,  riarhiiit;  a  iiia\imiim  at  alinill  thri'i'  tn  six  years  of  anc  aiiil 

••111  rapii|l\  ili'.liiiiiiL'-  tn  aliniit  t\Miit>.  afti'f  which  it  ih'i-lim's  iimcli  ninic 

''••■'•'!;..      The   ilirliii.'   in    th,    lariiil-  >.ais  .im's   imt    ( ''I'li   sli'ailil.\  .   Imw- 

I'l-.  for  at  till'  jti'iini!  jiist  pii'i'i'iliiin;  till'  niisi't  of  piiiii'it>    a  ili"iil.'il  iii- 
iH'O  l.i'i-iimis  i'\ii|i'iit.   imiii-atiiii.'  that   at   this  pi  riml   thi'   iin'taltnlism  nf 


B': 


w'l 

^   9 

-1  • 

■j   s 

IMt 

i 

l«.     ■  i 


'  I  i  if    ? 


Ik  i 


is:. 


512 


Ml'IAIKiMSM 


llic  '_'i-iiv\  iii'_'  iii-L'.iiiisiii  is  liciii<r  stiiinilMti'd.  I'"ciii:il<'s  liiivc  ;i  Iciucr  ciicij. 
iml|iiit  tliiiii  iiiiilcs.  unci  llic  >tiiiniliitiii'_'  iulliiciirc  of  piilM-rtN'  is  Irss  iiimiK'  ! 
ill  tliciii. 

In  rcMMiil  iiniiiliri's.  4(1  ( '.  pci-  si|iiaic  nn'tcr  of  suit'.ici'  pfi'  Ikpui-  is  t!. 
('iicri.'\'  (iiitpnl  III'  ndiMUiil  iiicn,  m  1.")  per  cent  dcvijitinn  l)('in>r  cnnsiiiii . ,] 
MS  ilc'cidc'illy  jiliiKiiiiiiil.  'I'lii'  iivi'i-jii.'!'  iMc'tjiliiilisMi  (if  f:it  :inil  tliin  suliJccN  - 
tiic  sMiiic.  lint  tliiif  111'  wiiiiii'n  is  (IS  pi'i-  rrnt  liiwiT  IliMii  llnit  (if  nii'ii.  '11 
lijisal  iMi'tiiliiilisni  III'  :i  j.'i'iinp  nt'  nun  .mil  wiiniiMi  iM'lwi'rn  flu-  ayes  nf  I'nii 
iind  tit'ty  was  \.'.\  piT  ii'nt  1  i''iiu  tlii'  avfi'aL'i-  I'uf  llif  laru'i'i"  L'l'onp  lii'tui"  m 
till-  au'i's  iif  t\\rnt\  and  fit'fy  :  and  tliat  nl'  a  jri'imp  lirlwi'i'ii  lil'ty  and  si\'-. 
vi'ai's  was  II.:!  pir  iM'iit  InwiT. 


M-   fiji 
\  V    ' 


!   3 


i  "  IT.   ' 


i;.  ;.   ! 

•;.f;       • 


i  .1. ,  i 


SI 


m 


1  "i 


Influence  of  Diseases 

Tin'  nicasiiii'inrnts  lia\f  Iii'cn  niadi'  liy  ttic  dii'cct  ini-tlind  wliiidi  lias  Jii-t 
lici'n   di'si-rilird.   Jml   sinci'   the   inurli   sinipliT   indiri'rl    inftlind    (  pau'i'  ."i."i  I 
yields  i'oini)aialili'  fcsnlts.  it  is  ln'ini.'  adiipt.d  I'm"  clinical  pui|)i>scs.     'I'll.  -' 
results  were  iilitainrd  li\    nnikinu'  i)aralli'l  dctciniinatinns  iif  cncriry  i'm' 
pnt    li\-   Iiiitli    Mu'lhiids,    in   disease  as  well    as   in   liealtti.      Sump  of  tlie  >■' 
siTvatiiins  that   have  lieeii   made  cm  the  ein't'try  mitpilt   in   varinils  disca^'  ~ 
aie  as  I'nlliiws:     In  \ify  sexei'c  cases  of  ( .niplilhnlniii    finih  r,  lirat   invidiii 
tiiiii  may  he  increased  liy  T.'i  per  cent  ovei'  the  normal:  in  seveiv  eases.  1.'. 
."id  per  cent.     The  warmth  of  the  skin  and  the  sweatins,'.  which  are  proni' 
nent    symptoms    of    this    disease,    are    therefore    aceonnled    for   liy    th(>    i' 
creased   elimination    of  lieat.   and    it    is  considered    possilile   that    the  otli'V 
sxmptoins  would  he  produced   in  an.N    normal   individind   were  his  metal" 
lisni  maintained  fiU'  moritlis  or  years  at  the  hiirh  level  which  it  occupies  in 
^'..itei'.       In    the    opposite    condition    of    iiii/.n  )h  iiitl,    the    eiieri;y    output    i^ 
markedly  i-cduced.  liut  rises  slinvl.\-  iIurinL'  treatment  with  tli\roid  e\trar'. 
or  much   moi'e  rapidh    with  the  very  active  thvroid  hormone  reeent.'x-  i-  ■ 
lated  li\-   Keiid.ill.       In  iliiihi  h  <  it   has  often  Keen  thou^'ht   that   the  r,ii.' ! 
emaciation  and  loss  nf  streUL'th  were  dependent    upon  an   excited  state  ■  ■' 
iiietaliolism.   or  a    useless   hurninir   up   of  the  ener^'x-    material.      The    m   ■' 
recent    work,   ho\\e\er,   cle.nl.x-    shows  that    this   is   not   the   ease,   the  Ii.i-  1 
iiiet.aliolism  as  ealenl.ited  jn  r  unit  of  liodv  suiface  heinir  within  the  liiiii'-. 
indicate  1  ;diii\e.      HuriiiL'  the  st,-ii-\  at 'on  treatment  the  enc'irv  output   ni-  ■ 
he  much   he'ow   the   noinial.      In    uneonmensated   cases  of  ( itril'ari  mil  •/ 
<  TV!  .   tl  ei'e   is   liii.i'e|veil    e'irri_'v   output.       In    l)i  r  ii'chnis   iliiiHiiil   the   metal 
lism   is  niirm.il.  altlor'j-h   in  '■evere  ca^es  there  may  le  an   increased  dem  r   ! 
for  oxyiri  n. 

Kvon  at  the  I'isk  of  repetition,  it  is  imiiortant  to  point  nut  that  in  :'"1 
?he-e  ii;-,.|v,  .^  th-'  :-..f:ry  ii,,(|-,,,(  1^  )],,  ..;,,iic.  ^v]i,-.f  lir-i'  iiieMsurcd  direct  iv  ■  ■■ 
h\'  the  indirect   meiimd    ''lut   to  he  deserihed. 


mi:t.\ii()I.ism 
THE  MATERIAL  BALANCE  OF  THE  BODY 


543 


We  iiiust  (lisfin-iuish  l)ft\vc(ii  the  l).iliiiicfs  of  flic  orj,'iinic  ,.  d  the  iii- 
aij.'iinic  foodsfull's.  From  a  study  of  the  former  we  shall  >;aiii  infoni;af ion 
ir^'ardiii",'  the  soiircis  of  the  I'licr^'v  lui.diiclioii  whose  lielia\i(.r  iiiidor 
\.iiiims  eonditioiis  we  have  just  studied.  From  a  study  of  the  iuorfraiiic 
laiaiM-e,  althou^di  we  shall  learn  nothing'  re^'anliiiv'  enerfry  exehan>;e — 
lor  sueh  sulistaiiees  eali  yield  no  ener-jy we  shall  lieeome  aecjuaitited 
uith  several  facts  of  cxtretTie  importMncc  in  the  maintenance  of  nutrition 
;iiid  j.'rowlh. 

Til  draw  mi  n  hnj.inir  shut  <if  iiri/inilr  iiilnl:,  and  oat  pat  re<iuire.s  an 
;Hiiirate  chemical  analysis  of  the  f 1  and  of  the  excreta  (uriiu>  and  ex- 
pired air). 

Methods  for  Measuring  Output 

The  principle  hy  which  the  output  is  measured  \v  ill  he  uiulerstood  hy 
1'  I'Tfinir  to  Fiir.  17(i.  from  which  it  will  he  seen  that  tlic  calorimeter 
1^  connected  with  a  close.l  system  of  tuhes  provided  with  au  air-ti^dit  ro- 


♦f)-*-   Water  to  obaorb   haftt 


I  >;-  !-■.  Ili.iKram  ut  At«al.  i  l.v.u-.iu  t  r,<|  ir.,li...,  ,  al,.,  ,m,  t,  , .  A~  ,!,.■  .■>i,:„.,l  „-,  .  i,,,  il„-  I)., 
!nt.,l    v..Im,..,-    ..t    :„■    sliniih-.      I  111-    >luii,k.iK,.    ,.    „„ii.  ;il,  ,|    l.v     ,|„.    „„.|,.,.    ;„„i    ,,    ...i  r  .m".".Iitii.- 

■v\.\    III    Biv.s    th,-    lO,;    ih.it    ,,|-    I.    llu-   «^,l,r    va|.n,.  "  '    ' 

.V  hlower  or  |iump  to  maintain  a  constant  current  of  air,  as  indicated  hy 
.■irrows.  Fi.llowiMir  !!■..•  air  stiram  as  it  leaves  the  cliainlicf.  wc  nol'r 
(Ic  tiilie  coniicctin^r  with  a  meter  to  indicate  chanjres  in  volume  of  the 


1 


J  j5 


I 


:. » I 


\ll  TM'.ciil-M 


.1     Ir 


'V    I 


;iir  ill  llii'  s\^liiii.  r.rvoinl  this  :iiii|  i|i,.  |>iiMi|i  is  ii  N|)fi'ially  ciiii.sl  nicl,  .| 
iiottic  ciiiiliiiiiiii';  iMiijcciiliiiIrd  1LS(>|.  tlii'ii  (iiic  coiitaiiiiin;  soda  lime,  iiinl 
lastlv  aiiollici-  ILSd,  li.ittlf.  'I'lif  lirst  ILS<>,  ImiIHo  alisorlis  all  tliu  walr 
\:i|>ui'  ciiiil.-niiiil  in  tlir  air  i-omin;:  riinii  llir  chamlicr;  the  soda  lime  imiti, 
.■ilisnrjjs  ilif  (  (  >_..  an, I  il,,.  m'ci.ihI  II,.SI>,  li.iltir  alisorhs  water  tiial  is  i>im 
(liircci   ill  till'  clii'iiiical    icactiuii    iii\(pi\ic|    in   till'  aiisi'-'jit  iuii  di'  the  I '<  •„  !._\ 

till'    Mida    li L'N'aOll   ■  (  ()_      llj  »      Na_('0    > .       |!\     w.'l'^liilif,'    llics."    ah 

Mir|itiiiii  liniilis  licliirc  and  al'Icr  an  animal  lias  liccii  I'or  suiiif  time  in  Pi. 
rhaiiilur,  till'  u.'i'.'lit  ot'  II  <>  and  nl'  ('(»_.  •.'ivcii  out  can  lie  dotfriuined.  Am 
ntlicr  side  liilif  lc;hK  In  all  i.\v;jvn  i->  liiiilrr,  till'  \al\('  III'  which  is  iiiaiii[> 
niatcd  Ml  as  l.i  i-aiiM'  nw.jcn  In  lie  discliaip'd  iiitu  the  svstcni  at  such  ,i 
rate  as  to  cmniK  n-alc  c\acll\    lor  that   Used  ii|i  liv  the  aliiiiial.  as  indicate. j 

\<\     llie    lichavior   of    ili|.    iiK'tii-.      The    ai Hit    oT   o\\  y:cn    l'e(|uirei|    is   <\r 

Icrniined  cillicr  li\  uciohiiiM  il,,.  ,,x\oin  c.\ltiider  liel'ore  and  after  the  .Hi 
scr\ation  ipr  lp.\  iim  „^uriiiy  llic  \o|niiic  oi'  o\;  .'ii  used  li,\  [lassiny;  it  tiiroii'jh 
a  carel'illl.v  caliliraled  and  \er\  seiisilixc  water  meter  insei'ted  on  the  si'l.- 
tiiiie  that  connects  the  ()  cylinder  willi  the  main  tuliiii'r  nf  the  systcMi, 
Since  muscular  acli\ii\  causes  pronounced  chaiii:es  in  the  rate  ot'  iii'' 
laliiilism.  means  arc  usually  taken  to  secure  i.'ra|ihic  records  of  aiiv  mo\.- 
meiitv  niaile  duriliii-  the  oliscrv  at  ion. 

'I'lie  ^rrouiiiy:   iiii|iortaiice    in   clinical    in\i'>tie-ations   of   iiicasiii-cmcnts  ..' 
the  rcs|iiiatory  exclianiri'  aiicj  tln'  neccssitv    for  lia\  inir  iiictiiods  that  are  :- 

siin|ile  as   is  ( sistent    witii    aceiifaey,   lia\('    led    to   the    inlroduction    "i 

sexcral  other  forms  of  a|i|>aratils,  of  which  those  of  I-",  (i.  jiencdiet  and  "l 
TiNsiit*  arc  the  most  iiii|iortaiit.  In  tlie  former  a  tiuhtly  liltinu'  ma^  , 
a|i|ilied  o\i'r  the  nose  and  mouth  is  eonnected,  liy  a  simrt  T-pieee,  with 
the  same  tuliinir  as  that  used  in  the  res|iiratioii  calorimeter.  The  patient 
lliiis  liri'athes  in  and  out  (d"  tiie  air  strcain  that  is  passinir  alon^'  the  tuhiiij 
\i  itiiout  any  of  I  lie  ohstriiel  ion  e\|iei'ieiieed  w  iien  the  IireMthini,'  has  to  ' 
])erformed  ihroiiah  \al\es,  as  in  the  older  Ziint/ ^  forms  of  )i(irtah  . 
res|iii-ator>  apparatus.  It  is  particularly  for  studies  on  man  that  tl  '^ 
apparatus  has  lieeii  <le\  iseil.  The  Tissot  and  DoiiciJas  nietlio<ls  are  sine  ii 
in  Kijrs.  17!l  and  1mi.» 

To  complete  the  invest  ii^at  ion.  it   is  necessary  that  the  urine  and   I'c    - 

1 illeeted  and  the  nitroiren  excretion  nieasiired.     When  the  respirai' 

excreta  are  measured  o\er  a  coiisideralile  perioc]  of  time,  as  in  the  lai  :■• 
caloiimeter,  the  urine  is  enllected  for  tlo'  same  period,  hut  when  shot'  '• 
respiratory  measurements  are  made,  the  urine  of  the  twentyd'oiir  ho  • 
is  Usually  taken 

Principles  Involved  in  Calculating  the  Results.     Provided  with  the  . 

al\  ses   fiKiiiNlieil   !i\    !i;r  a!;o\  i-   nirthnds.    v,  e   piiici'cd    to   ascerfaill   flic   ti'    .' 


.I...I    v,.;:    I,.     I..  ,,  .1    ,|.  -.Ill,, 


MITAnoi.ISM 

545 

.-imount 

s   of 

niti'op'ii 

iimI   ( 

•ill- 

(Oil    excreted 

and   t 

o   ea 

eulate 

ffom 

tlie 

Kiiiiwn 

CIIIll 

iDsitii 

III     of 

prut 

■ill 

IliiU     IllUeli     1 

nitciii 

must 

liave 

UIli 

ei 

Kono 

!iu'tal»i 

ism. 

Wo 

then 

Cllllll 

Ult( 

'    lliiw    Illllell 

•ariiim 

this 

quantity  of 

pro- 

ti'iii   Wdllld  . 

KMMJllIlt     lol' 

,  iiiiil 

« 

•   (ledllet    ihis 

from  the  ti 

tal  ear 

lioil 

oxcre- 

(III.  The  remainder  of  carlion  iiiii>t  liave  come  from  tlie  metabolism  of 
its  and  <'arlMihydrates,  and  althoniih  we  eaii  not  tell  exactly  which,  yet 
e  can  arrive  at  a  clcse  apiiroximation  hy  oliscrviii<r  the  resfiiratory  (|Uo- 
I'lit  (  K.  i-i).  wliicli  is  the  ratio  of  the  volume  of  caihoii  dioxide  exhaled 

1  that  of  oxy;:eii  I'etaiiied  liy  the  liody  ill  a  u'iveii  time.  i.  '•■•— tt  "   .    15y  "h- 

■rviii^r   this   (|iiotie!it,     therefore,     we    can     approximately   deteruiiiie    the 


iirce   I  roll!   u 


liicli   the  noiuHotciii  earlion-excretion   is  derived. 


Ilaviiif;  in  the  al)o\(-  iiianner  coiiipiited  how  much  of  each  of  the  proxi- 
late   principles   has   unileijrMiie   iiii'taliolisiii.   we   next   proceed   to  compare 


itake  and  output  wil'    a  view  to  tindiii'r  whether  tlie 


re  IS  an  eipii 


liliriui 


ImIwccii  the  tw( 


ether  retention  or  loss  is  oci-urrili<'. 


It  iiiav  sirve  to  make  clcai'  the  mcthocls  liy  which  these  calculations  are 


to  stud\   the  t'olhiM  ilitr  exaiiiple: 


F.  XII  lit  I'll-  iif  II   Ml  liihiilisin   lull  sliiiiiliiiii.  '  At    IS  ili'-iiid   tu  ki 


iliilliri    ii  iliit  I'lm- 


lis  pintiiii.  ."ill  ^riiiiis  t';il.  nil. I  ."ii'O  L;r;iiiis  iiulioliv.iriitr  i-^  siilliiiriil    I'l.r 


i^iiiiiii;;  1_.>  ^rjo 

1'  111  (liiinii  11  iii.iilcTuli'  iiiiiiiuiit  III'  wink. 


I  NT  A  K  I'. 


fail.. 


Xili 


Trcitein. 

Ciolii.livdr.'ilr 

Fiit. 


CM.. 


2".")il.O 

4i;.-).() 


T.iUil, 


I'.oo  ••m. 


20  L'ln. 


OlTITT 

('Mil... II 

Nitn.^rn 

Ill   iiriiic, 

11  mil.  ( 111 

..')  ■  0.117) 

1(1..")   ;;iii. 

III    r.'.'i's. 

.") 

J.ll 

Ill   til.'  l.r.':illi. 

::."ii 

— 

Tiital, 


•270    j;lll. 


17..")   am. 


liiliiiiiiil  ill  liii'lit.     .'.0  uiii.  (•:irl...ii  :ti..l  1'..")  ::'ii.  nil  ii.^.'ii.  Tliis  ainouiit  ..(' iii;  r..;;i'ii  ri'prc- 

-•  Ills  -..*)■.  11.1*.") —- l.").ll  ^111.  piiiliiii  <']■  7.")  ;iiii.  iiiiisi-h-.     Now,  tlii(  aiiumut  i.it'  pL.tciii  will 

■.!■.), lilt    f.ir   s.l.*.")   ^Mi.    rait.nn;   sii   that   .'Hi     s.U*.") -- li  1 .7.')    liiii.    cailniii    ii'|.ii's.'iil>^   1*1.7.")  • 

'.:l^"S..",  ;;lil.  fill.     On   this  .lirt,  tlieli't'urc.  th.>   slll.jrct    li'tiiilis   ill    his   tisMl.'s    l.j.ti   ylii. 

■  i.li'iii  anil  -S..'l  ;;iii.  fat  prr  .lirni. 

I-'urnished  with  these  data  we  ma\    now  jiroceed  to  coininite  how  much 

'  iMi'}j:y  must  lia\('  hecii  lihciatcd  in  the  liody. 

'I'll  express  the  almvc  result  ill  terms  of  eiicifry  lilierateil.  we  know  that 

"-7.5  ('.  wore  supplied  and  that  all  these  have  heoii  used  except  15. (ix 

i.l     r>4  retained  as  protein,  and  2S.;!  ■■  !).:!     2(!:!.2  retained  as  fat:  or  in 

•  I'l  ■'<2~.-2  ('.     We  lind,  thiMr.f.ir.',  that  :!ni'7.5  —  li'JT.-J  == -JJOd  ( '.  jiav"  Ik'cii 

:i  i|uired. 


I'        II 


i. 


1.*  '  ■ 


iJ:' 


r 


•iti 


Ml  TAIl(Pl,l>M 


:i.i!- 


Tliis  is  called  tlic  iiicthiMJ  at  iinllrii  I  i  ulnri'in  Ir/i,  iiiid  it  liiis  Ixm'ii  clcijilv 
fNtiil)lislu'(l  liy  iniiiicriiiis  nhscrvatiuiis  that  tlir  irsiilts  asrri-o  exactly  uiili 
tluise  sceuii'ij  liy  the  iiiethod  of  ilinti  i  nlnrlnii  trii  ileseriliod  aliove.  l-'.^i- 
most  iMU'imscx  the  iiiilii-ect  iiiethcid  is  (|iiite  satisfactory,  ami  il  is  es|i.'- 
ciall\  \aliial>le  in  cases  in  wliicli  thei'c  are  considcfalile  and  siiiMi  n 
chaii-fci  in  liody  teiniieiatnrc.  That  the  results  l)\  the  lw<i  methods  shoiil.l 
a'.'i'ci'  shows  clearly  that  the  law  of  the  conservation  of  enerjry  must  apph 
in  tlie  aninnd  hody.  for  il  is  exident  that  if  any  eneryy  were  derived  fii m 
outside  the  hody  other  than  that  taken  with  the  food,  tin-  results  li\  l|w 
direct   metliod  would  lie  liiL'her  than  those  liy  the  indirect. 


rilAI'TKIf  I.XI 

TIII-:  (AIMJnN   |'..\|,\.\(  !•: 

Before   piiicooilinj.'  to  discuss  tli(-  sp.ciMl    inctjilioliMn   of   iiiutcins.    I'iits 

iiiul  cjiriKiliyilijitcs.   it   will   l.c  M(]v;iii1ii>;i'i>iK   to  coiisidir  liriclly  s <ri'ii- 

iiiil  fiicts  (•oncfniiiit;-  the  cxi-rction  oi'  .MrlMni  .lioxi.lc  and  the  intake  oi' 
'i\y<ivn.  In  the  first  plare.  il  is  iiripoitant  to  note  that  tlie  i.iI.hI  of  th.' 
conihiistion  pideess  in  the  animal  hodv  is  |pfo|(o!ti()iial  lo  the  amount  of 
oxyiren  ahsoHied  and  of  carlion  dioxide   produccil.  whefeas  the   imhin    of 

tlie  eoinlillstion  is  indicated  liy  the  ratio  existin;:  lietween  the  ; unts  of 

carhoii  dioxide  expire.l  and  of  <ixy<;cn   retained   in  the  liody.     An   invesli- 
■ation  of  tlie  earhon   halanee.   in   other  words,   is   partly   ipiantitativc   and 
partly    <|iialitative  -<|Uaiititative    in    the   sense   thai    it    indicat.'s    how    in 
tensely  the  liody  furnaces  are  Imrinnfr.  and  ipialitative   in   the  sense  that 
il  tells  lis  what  sort  of  material  is  lieiny:  liurned  at  the-  time. 

THE  RESPIRATORY  QUOTIENT 

Influence  of  Diet.--  The  res|)irator,\  (piolient  is  determined  liy  com- 
parison of  the  volume  of  carlion  dioxide  expired  with  the  volnme  of  ox.v- 
LTi'n  meanwhile  retained  in  the  liody  or.  as  a  forninia. 

\'ol.  Ct »..  expircil 

\'ol.      <».   ri'tained 
l-'or  the  siike  of  brevity  the  respiratorx  cpiotient  is  oI'icm  u  i  iien  H.  i^.    Thai 
il  servos  as  an  indicator  of  the  kind  of  eondmstion  occurriiii;  will  lie  evi- 
dent from  the  I'ollow  inu  e(|nations: 

\.  r,i,hi,hiiilnit,  :   f,  II,  (),     i!0  —  i;((>      i;ii() 

( ncxtiiKc,') 

CO        c, 


RQ. 


(I 


1. 


•-'.  Flit: 


."..  Trotfiii  ; 


•  Mr  ((■   II   ()         sdO  —"CO      .-L'llO 
(Olcin.^ 

'  (),  so 

(  .  Tt.,,x„n  ,s  ;■  7:0  — ti:;r()     .'.siio    iifOiN'ir  V    sd, 

I  l')in|iiri('al  t'liiinulii  fm 
!illiiniiiii   (  I.iiticiUiiliii  1.1 


w; 


r.4H 


MITAIiOI.ISM 


I'i.ll 


II 


^.i 


I.    ('ini  14  isinn   itf  flit   iiitit  I'llihuhinhilfi  : 

-<',II  (<',J|    ()  I,  .  titO      -  lllC,  ll,<),       IsCO       vIl.O 
(Olcill.) 

'  ( )^  C.  I 

."l.    Cnillt  i:^i",i    >>f   ritt  hnlilfittillf   illtll   II    htijiil    fill: 

i:;(',n,(),.  —  ('..ii,./),, '  uiico  ■■  -nwy  o. 

I  (>h'"^traii'|iiilriiilifi. ) 

Tiikiiiu'  ciirliiitiv  (li'iitfs  first,  tlic  [.'('iici'ill  forniiilii  iiwiy  !u'  written  ('II  <> 
friiiii  uliicli  il  is  iihiiii  (li.it.  til  i>\ii|i/.i'  llic  innlfcill",  (i\\t.'('li  will  lir  i. 
iliiircil  ti)  i-ciiiiirnir  willi  tlir  (•.•irliuii  alnnc.  iicrnrdin;;  tn  flic  «'i|iiMtiii'i 
('II. <•  U  (  O  ,  II  o.  Ill  (iiliiM- wiinls,  till' xkIuiiii' of  ( iirhdii  (lidsiili'  pre. 
iluiTil  liy  till'  (Miiiilillstiiill  will  l)f  cxiirfly  i'i|U.ll  to  the  vnlllllic  of  dxv;.'!'! 
llMil  ill  fllis  iPi'iirrss.  ill  (iliciliilU'C  tn  flic  wcllklliiWll  (Jiis  liiw  fllilt  I'lpii 
IiinlciMlliir  i|ll;ilititii's  nt'  ililTii'i'lit  irMsi-s  i:<m'U|i>'  tin-  same  viilililli'.  'I'h. 
n'S|iir,itiii'y  i|iiiitii'iif  is  tlit'ict'iU'c  unify  (  Iv|niiliiiii  !■.  Wifli  f.ifs  iiiul  pi" 
fi'iris.  liiiucvi'i'.  til'  '.'ciii'riil  fiii'imilii  must  In-  written  ('II,  M),  iiuliciil  iirj 
flicrcl'iirc  tliJif  \'ifV  its  cipiiiplcti'  (ixiil.itidii  flic  iiiolcciilc  must  lie  siipplii.! 
with  nxxiri'ii  ill  snlliciciif  iiiiioiinf  fn  i-niiiliinc  not  only  witli  :ill  ol'  flic  imi 
liiili.  hut  .ilsii  with  siiiiic  III'  tlic  li>  ilrujrcii.   fiil'iiiiii^r  wnfcr:  so  fli:if   flic  v^l 

nil f  ('(t.   piimIiiitiI   will   Ic  /(.\\  fliiin   tlic   vdIimiic  uI'  nxytrcn   I'ctjiiiicl 

:iiiil  flic  rrspir;ifiir>-  iumticiif  will  lie  less  tliiin  unify.  As  ii  iii.-ifter  oj'  |";i.-i. 
lis  tlie  mImixc  ei|UMtiniis  slidw  {'1  ;iiii|  •'!  * .  file  rcspirafiiiy  ipidtieiif  I'nr  I'.il'- 
;ini|  prnteiiis  lies  sipiiicw  licic  'ictwccii  (1.7  .Miil  D  S,  lieiiiir  llslliilly  iie;iii-i 
(I  7  in  fl nse  111'  I'iits,  iiinl  iicMler  tn  d.S  in  tlic  c;ise  nt'  |)iiitciiis. 

Tluit  tlic  eiiiKlitiiiiis  liypiitliee:iteil  ill  tile  ci jll:itiiins  exist  ill  the  iiiiiiii.il 
lidily  (liiiinir  tile  cdiiilnisliuii  nt'  tlic  I'limlstun's  ciin  cisily  lie  shown  liy  nli 
sci'vinir  flic  respli'iifiiiy  i|iiiitienf  nf  iiiiiiiials  nn  ililVi'rcut  diets.  An  le'i' 
Ipiviiriius  Miiiiiial.  siH'li  as  a  raliliit.  when  if  is  well  fed  <_'ives  iiivariaM>  .1 
respiratory  ipintient  of  alioiil  1.  wliei'cas  a  strictly  carnivorous  aiiiinal. 
such  as  flic  cat.  trives  a  respiratory  ipinficiit  of  alioiif  0.7.  Hvcn  liini' 
striUiiiL'  jierliaps  is  tlic  eoiiipai'ison  ol'  flic  respiratory  ipioficiifs  in  ai' 
licl'liivoiiiiis  animal  while  it  is  well  fed  and  after  it  has  hccii  starved  for  . 
(hiy  or  two.  Ill  'he  hitter  case  the  respiratory  ipioficiif  will  fall  fn  a  ln\' 
level  liccausi'.  liy  sfar\  atinli,  the  animal  has  liccii  <'nnipclled  to  ehaliL'c  it- 
enniliiisf ion  material  from  the  carholiydratc  of  its  food  to  the  protein  ai;  ' 
fat  of  its  own  tissues. 

.\s   already    explained    ( paj.'e   .■■|4."i).    if    is   from    the    respiiMfoiy  (pioti'i  ' 
fliat    w  c  arc  cnalilcd  to  fell   ^\llaf   ))roiiortioiis  of   fat   and  carhohydral' 
respcctixcly.  are   uiidcriroiiiir  mctaliolism.      A    useful    falile   sliowiii<r   tl 
jicrccntairc  of  ealoiics  piii(|ncei|  l»y  each  of  these  fooilsfnlVs,  iiffcr  allow 
iiif.'  t'or  iirofcin.  is  irivcn  liy  (Jrahaiii  Lu>k   (sec  pa^'e  .")(i.")). 


Tin;  cAiiiiov  ii.M.ANci-: 


549 


Influence  of  Metabolism.-  Ai)iiit  fiom  .lict,  tlio  rcsiiimlory  (niotiont 
iii.iy  often  lie  iilt(>rc<l  l)y  <-Ii;niui>s  in  llic  nicliilxilic  lijiliits  (if  the  aiiitnal. 
The  <•  lire  iiir.st  ciiiisiiiciHuiHly  cxliiliiti'd  in  flip  ciisc  (if  liilicrtiatiiif; 
.iiiiiiinls.  In  the  aiitimni  iiKintlis.  wlicn  llic  animal  is  patMisj  v(iraci(iiisly 
■  if  all  kinds  (if  catlKiliydralf  fddd  aiul  dc|)(isitiii^  larw  fpinntitios  of 
.idi|)(is('  tissnc  in  liis  liody.  the  rcspiratdry  ipKiticnt  may  lie  cunsidcralily 
■jicatcr  llian  nnity,  indicatinir  tlicrcfdrc  citlnr  that  relatively  mdio 
i'.iil)on  dioxide  is  lieinLr  diseliarced  or"  less  iixNU'cn  fetaine(l.  As  a  matter 
df  fact,  it  can  easily  lie  shown  that  it  is  the  fdrmoi'  of  the  eauscs  that 
is  iTsponsihle  for  the  hiirlier  (lUotieiit,  the  explanation  for  the  inei'eased 
lirodliction  of  ('( l^,  heint:  that,  as  the  cailiohydratc  elian<_'es  into  fat,  the 
relative  excess  df  carlion  in  the  fdinier  is  i.'iit  rid  df  as  CO,,  as  indicated 
ill  K((nation  .").  On  the  other  hand,  if  llie  animal  is  examined  uhile  in 
liis  winter  sleep,  it  >\  ill  lie  t'onnd  that  the  res|)irat(iry  <|iiofient  is  now 
I Atrenn'ly  low,  often  not  nioie  than  (I.:?  to  (1.4.  which  may  ]>■•  intcipreted 
..s  indic.-iliiiu'  either  an  exeessJM'  alisoiption  of  oxy-ren  or  ji  markedly 
'lecreased  excretion  of  carlmn  dioxide.     .\s  ;i  matter  of  fact,  there  is  a 


t  (1  niinnlion  in  hoth  the  excretidn  (if  c.-irtion  didxide  and  the  intak 


lremel\ 


le  ;inimal   is  extremelv 


lei;rt 


less  (leifref 


urea 

I'f  O..,  liecanse  the  whdlc  mctalidlic  activity  of  tlr 

dcpicsscd,  lint  this  (liminntion  alVccts  the  oxyycn  to  a  much 

iiidicatini;   therefure   a    relative    increase    in    the   oxyL'en    retention.      The 

■  xplanation  is  that  the  oxyijen  is  lieintr  used  in  the  chemical  proeoss  in- 

\  dived  in  the  conxcrsion  of  the  fat  hack  into  carhohydrato. 

Whatever  may   lie  the  relationship  lietwccn   fat   and  earliohydrate   in 
llie   nonliilicrnatin!,'   aninial,   then     is    no   donht     that     durin<;    hilicrna- 


dii,  hefoic   the   fjit   stores  ai-e  hnrned,    fat    i 


dsely  I'cl.ited  td  c;irli(ih.\  (]r;it,'s.  tlh 


itcd 


s  cdnverte(l   into  somethiit!; 
('(piation  I'm-  the  jtroccss  lieini;  reii- 


iis   "iven   a 


hove   (  \o.   4"!. 


In  man  and  the  higher  maiuinali.i.  the  only  condilidii  apart    from  diet 
liich  can   afl'ect   the  iiatl 


e  ot    the   conilinsllon    p|-occss   is  dise;ise 
cs    (  pa^'c   (iTSt    the  orL'.'inism    loses   the    pdwer   (if   li 


th 


II   tdtal   dialiet 
'  nilidhydratc,  so  that  \\hatcver  the  diet  may  he.  the  I'esjiirntdry  (pKitient 

^  very  low,  never  hi«:her  tlian  that  representing'  comlitistion  of  fat  and 
I'i'dtcin.  It  lias  liceii  claime(l  hy  certain  invest ii,'ators  that  in  diahetes 
■lie  respiratory  (piotieiit  may  f;dl  cunsidcralily  helow  0.7,  indicatiiii.',  as 

11  hihcrnatinv'  animals,  that  tat  is  l)eiii>:  converted  into  earliohydrate. 
llie   most    recent    ;ind   carcfnily   c(inlrdlle(l   oliscrvatiims,   however,   denv 

iiis  claim,  and  for  the  present  we  must  assume  that  in  the  liody  of  man 
■  :i  is  lid)  cdn\(Ti(d  into  carlidhydr.iTc  i  se,-  p.iirc  Ilii4i.      In  nuiiicrous  ulticr 


iiscjiscs   investiKate(l   hy    |)ii   linis  and   othe 
'  lie  comhiist 


s'    no  (pialitative  chaiifie  in 


comhiistion  iirocesscs  in  man  has  hec'i  hroiight  to  lii[;ht. 


■Vl<l  Ml  •|Alll)|.l»M 

THE  MAGNITUDE  OF  THE  RESPIRATORY  EXCHANGE 

It  is  c\ji|clil  lli;il  llic  iiiiKiiilit  111'  cjiilioli  i|iii\ii|r  cxpil'cil  illid  ii|'  m\\ 
U'cll  l.t.iiiinl  will  lie  |iri<|iiirtiii|i;il  I"  llic  cllcisiV  lihcliltioll  jli  the  iiniiii;!' 
liiiily.  \-',\i\]  ill  ihr  lisl,  III'  ri'|ii'tiliiiii  it  nIhhiIiI  lie  iintnl  tliiit  tli- 
(•liriuy  r\i-li;iinic  rjiii  lie  \ii\  iin-ili;it<l\  nili-iiliiti'il  I'lnm  tin-  icslilt  nl' 
liic  iiijilrii.il  li;iliiiMi'  -IiitI  iinliii'i-t  ciiliiiiiiii'liy,  iis  it  is  called  (piiL'i 
.>•)■_''.  Oil  ,'icciilliil  111  till'  iiiiN|i.ll;ili\c  silii|i|icil\  111'  liicjisUliM;.'  llic  c.llliiii 
iliii\iili'  iiut|iul   iiiiij  uNv.'cii   inliikc,  it   is  n;iliiii;l  lliiit   iiiiiiiy  nf  tlic  iilisii 

Miliiiiis  tlijil    have  licni  niiiil i  incrcy    iii'iiiiiicliiiii    in   tlic  iiliiiiiiil   Imil 

i|c|iciiil  nil  the  use  111'  tliis  iiictliiiij.  Jiisiiliciitiiiii  I'm-  wliidi  is  riniiiil  in  lie 
i-iini|ilclc  Murcrniciit  liiiwccii  liic  icsiilis  nl'  ijircct  iiiid  indirect  calinini 
dry   ill  a   ^rcal    \aiicly  ul'  diseases  and  cnnditiiiiis  in   niaii   (|lii  Uiiis'  i,' 

In    tlic    first    |>laec.    it    is    intci-cslini.'    t iii|iai-c    llic    rcspiratnry    es 

clialmcs   111'  liirrcreiil    animals   ciiininited    jiei-   kiln    lpiiil\     wciiilit.      This    i- 
sliiiw  II   ill  tile   I'lilliiH  ill'.'  lalile. 


!    't 


I'M 


II   « 


m 


-1 

■ 

!■ 

1  - 

} 

[U 

(^ 

I       iiXVIll-N    Ml  I  AiaiO.N      IHilMlii:  I 

will, HI         1,-iililll  11  I'l  l:    Kll.ii         lilSlllAKiait       !       Mil,.    III.  TK.MrKnx- 


wii  inn  K 

l.M. 


ri  K   Kii.o 

\  -II  inn  i; 

l.Af, 


I'iil.l  i-iiikii  i       ii.L'.-i 

Inifhihi,,  I 


rniKiit 
.Mi; 


i:.iii.i,.  fi..-    ; 

11. m;;; 

(  II. •.'...•.. 

11. mill 

'  ■:ii,7ii  .•..-.  • 

ll.lill 

I.-.    i;i' 

II.  111.*) 

ii.ii.';i 

II. 7> 



i7.:.liM-.i 

1  .")7    ~  IM'     ) 

■    ..    .  . 

.(...* 

CiiiiiniiiM   lii'ii 

iL'sli 

1.li.-,S 
7  III. MM 

I..'::.'7 
1  ii7.";  .■..-.  1 

ii.'.il 

111" 

l'i-r..ll 

L'-'l«  ilsli 



:i.'j.':i! 

-^ 



Spurinw 

1 

'-'- 

!i..")!t."i 

1  II7III  IM'.") 

lll.tHL' 

11.7!' 

IS" 

l/.ii. 'i-i 

IK                       J 

lloS,|lllll 
(ilin.lllMl 

- 

.i..",s;i  II,  |s.-, 

-- 

— 

Sl,....|,                  1 

lili.tlllll 

Il.l'.lll 

.:;i::i-..'.i 

im!7I 

(.'Ml  .MM 

ii.fi'.i 

ir.' 

I),,-             1 

1 

Ii:;!:: 

l.:;ii:! 

I'.UI    .MM 

i.:;-.'.'i 

iil7l<M'.-| 

(1.71 

l.V 

«';,f 

Uliil 

1  ..-..-.Ii 

t.:;!i7 

(1.7.-) 

n   .10 

:;iil7 

(  ;i  17  .'..'.  ' 

171II.M.-1 

■ ' 

ii.lll.-, 

ll.7lirt 

II. si; 

U'.'.lt  ' 

.  l.-.n  .-..■.  ^ 

.:;mi.m-.i 

lllll.l.it 

1 1;'. 

l.iilJ 

i..;."ii 

11. '.17 

I^     iJn 

lillilllll    I'i^ 

Ill  '.1 

LIT- 

l,7.-.s 

11, si: 

|{;it    Mvliii. 

*ii.."> 

:;..-,  Is 
.  i7^:i...M 

— 

1 

.M.iii 

•_'.-, 

s.t 



17° 

Mm. 

I'l'. "11 

n.ll!i- 

(i.r'.L'T 

— 

(  Mii.liiic.l    fiiiiii   lVmIiioy.)i: 

"     ]•"■'' il >'"•    '""     ■''•■ !-■■•■>     1 Ill       iii.lllo.U    ,,f    .■,il,,K- 

'll.''l.     Ill     II"     •  liin.  ,     .    .1m|.-.i     .,11    III,     -;il.i,.  I     ^^lll    l„     l.ucl    lirtnil.llK    "11    ).;1K'-    .=  .^1. 


rill    I MdiiiN   iiAi..\N(  i: 


il 


S<'\«'i-iil   t'iicldis  iipcriilc  til  ('\|»liiiii  tlifsc  (lilTni'iircs,  iiiid  nf  tlicso  flic 
ihIIdw  inir  air  of  iiiipiiitiiin-c: 


I.  The  Body  Temperature.     I 


liiTfiisr    ill    IhiiIv    ti'lii|i<'l'iitlirt'   t'litllils   iil- 
I'ciisod    ('(11111111^111111.      This   cMiliiiiis   wliy    llic    iiictiiliiiliKiii    (if  ii    Iiird    is 


II  k 


IKIWII,    tiK 


tll< 


irrciiItT  fluiii  that  (if  »  iiiaiiiiiiiil        the  same  size.  f(ir.  as  i: 

iciiilionitiirc  (if  a  liinl  is  twd  (ir  tin Icu'rccs  (•ciitis.'rado  ali(iv(>  tlint  (if 

iillicr  aiiiiiials.     Hisc  in  limiy  tciiipciatiiic  also  explains,  in  jiart  at  ]vasi. 
the  iiicrc.isfd  iiictalidliMii  (ilisi'ivcd  in   fcNcr. 
•J   The  Temperature  of  the  Environment.     In  ciiisidciiiii;  this  \\p  must 

•  listinifiiish   lictwccn    tl tVcct    prmliiiM-d    (in    wanii-lildddcd    and   on   cnld- 

I'i led  animals.     Since  tlie  Imdy  teiiiperatiire  (if  a  edld-lilodded  aiiitiinl 


llv    (11 


le  (ir  t\\(i  decrees  ( 'entii;i'a( 


le  al 


idvc  that  (if  its  eiivii'diiinent.  it 


I'ulldws  fliat  the  iiietalidlic  activity  will  lie  direefly  pniportidiial  to  the 
li  iiilicratiire  of  tlic  latter.  In  a  w  anii-lildodcd  animal,  (ni  the  other  iiaiid, 
the   liddy   temperature    remains   cuiistaiit    whatever   eliaii(;es    may   occur 


that  of  the  eiiviniiimciit,  this 


constancy   of  liddy  temperature  lieii 


licpeiideiit  (111  the  fact  that   the  intensity  (if  tin mliiistidii  jinicesses  is 

iiixcrsely  proiKirtidiia!   to   the   iliiiir  eft'ect   of  the  atmosphere.      Thus, 

Mippose  the  external  teiiiperMtiirc  sliduld  fall,  then  the  hiss  (if  heat  from 
ilie  liody  will  tend  to  licconie  tricater.  and  to  iiiaiiitain  tlie  liody  teiiiporn- 
tiire  at  a  constaiil  level,  the  lioily  t'liniaccs  iiiiist  liiirii  more  briskly,  with 
the  result  that  an  increased  excretion  of  carlioii  dioxide  and  intake  nf 
"ixyiren  will  occur. 

This  intliience  of  the  Mirroiindiii'i  atiiiosphcrc  on  the  niefaliolic  activ- 
il,\  of  waniililooded  anil  nls  has.  as  already  iiointed  out.  lieeii  used  liy 
several  investijiators  lo  explain  the  greater  coinliustioii  iicr  kilo  body 
\\cii;ht  of  small  as  compared  with  larce  animals.  The  ar<.'iimeiit  is  that, 
viiice  the  surface  of  •.mall  animals  relatively  to  their  mass  is  iinieh  greater 

iliiiii  in  larire  animals,  tin oliiiH;  nf  the  small  animals  will  l-e  iirojiortion- 

.iiciy  <;re;itcr.    Tiic  rclat icmsliip  lictwccn  surface  and  mass  is  shown  liv  tak- 


•tl 


icf;   th 


e   lii.-iss  II 


I   the  twi 


i-lll 


o  clilies  IS 


ma  two  clilies  and  piittiiiL'  tliciii  1i 
I'lUal  to  doiilile  that  of  cither  iiilic,  whereas  the  siirt'ace  is  less  than 
'l.iiilile.  since  two  jispccts  nf  tlie  ciilies  have  Iiceii  liiou}.dit  totrelher.  To 
]'iii\e  the  ciinteiiliiin,  the  resiiiratnry  excliainrc  has  liecii  en;.  >uted  per 
~'|iiare  meter  of  snilai  c  instead  uf  )icr  kiln  limly  '\eiirlit,  with  the  result 


'ir.ti  a   \ crv  el 


nsc    en 


ricspnliilcncc    111    the    III 


'talinl 


ism  of  different 


aniiiinls 


.IS  lieeii  iilisciN  ('(!  ;  lull  this  i|iicstinii  has  .-ilreaily  liccii  discussed,  and  we 
■  in  know  that  the  law  of  cn.ilin;;  can  not  lie  the  mily  one  that  detciiiiiiics 
\leiit    of  the   rcspir.itmy   cNclian^rc    (sec    pa<.'c   .'141). 

3.  Muscular  Exercise.  This  ii.is  a  most  inipoitant  iiitliicnci'  on  the  ex- 
'.iiiire  and  it   is  part hiilarU    in  iniiiicetinn  with  i;   that  studies  in  carlioii- 

'i\iile  niitpnt  and  ow^cii  intake  have  liecn  of  irreat   practical  value,  p.ir- 


if 


'4 


I,,'' 


:t,i^ 


ji- 


'il 


1^- 


u 


flTW 


MITAIIdl.ISM 


tirlllllllv   xvlicil  till'  ilivi'stiu'llf iiUlH  )iir  lliuli'llill<rii  dm    mi'll  ilniliu'  nlilinai  . 
1,\|n's    (if    iiillsriiliil-   cXrl-riM',    Mli'li    iis    wiilkiliu'    "l'    rliliiliiiii.'.       It    is    tfii 
lllflt    till'   ililtlli'lirr    dl'   lilllMMllar   I'MTcisc   nil    the   clii'fL'y    mi'tllliiplislii    MK! 
(iImi  III'  stiidicil  l>\    liii\iiiL'  i\  pcisiPii  ill  till'  I'iiioiiiiicli'f  del  exercises  ini  a  i 
erjriiiiieler.  liiit  tlie  lesiilts  lliiis  olitiiineil  iiie  in  iiijuiy  \Miys  nut  iieiiily  v, 
\  iilllillile    !is   those    wiiieli    ejill    lie    seemed    liy    ciliset\  iiiir    tile    ivspir.itni 
exeliiiiiL'e  nl'  persuiis  diiiiijr  mdiiiiiry   types  nf  iniiseiilnr  exereise   in    tl 
open.     The  rnilnw  ini,'  t.ilile  of  iilisei\  ;itiiiiis  nii  lioises  is  ol'  interest   in  tin 
ciiiineetiiiii. 


eiiMllTlilS 

■Ml!  fM'IKUl 

cAiaii'N  nmxiiiK 

iiXMa.N  Misdiauii 

, 

IS  I.ITI  i:s 

iPisiiiMuaii  i\ 

l\-  I.ITKKS  I'll; 

II 

I'l  i;  MiM  Ti; 

I.ITK.IIS  I'Ki; 

MIX!  TK 

MINI  TK 

i.irs 

Hist 
Wnik 

177 

l.tWII 
I.Tfiti 

11 

■rriif 

Vi'I.'l 

r.aiii 

s.iiii;; 

II 

It    uill    he  iiliseived   Unit    the   iiiet.iliiilisiii    iiiereases  extnundinaiily    I'" 
even  ii  niodeiate  deu'iee  nj'  uiiik.  lint  that  at  the  same  time  the  iTsi)iiatiii 
ipiiitieiit    remains  eunstant.      j-'nuii   tiliser\  iilimis  nn    the   fespiratory   i^ 
ehaiiL'e  >il'  •Mirkini;  men  and  animals,  extremely  imp.iHant  faets  ediiceii, 
iiii:  the   enieieni-y   (if   niuseiilar   wmU    ha\e    heeii   secured.      The    fnrm    "i 
respiiatiii-y   apparatus   iZniit/   »r    Dmifrlasi    employed     for    tliis     piirpn-' 
iriiist    he  eapahic  of  liciiiu;  strapped  on   the   man's  liaeU    without    caiisin 
any  eniliairassment   to  his  hodily  movcnniits.     I'.y  a  comiiarison  of  tl  ■ 
respiratory  exehantre  with  the  amount  of  work  done,  the  eflicieiicy  of  lip 
work   can   rea<liiy  lie  determined.      It  has  liecii   foiiiid,   for  example,  tha' 
the  enicieney  is  iniich   frreater  after  the  man  or  aninial   has  triit  into  tli' 
su  iiiy:  of  the  uiirk,  his  eneriry  expenditure  jier  unit  of  work  heiii;r  muc! 
U'l-eater   dniiiifr   the    liist    half   hoiii's   work    in    the    morninj:   than    it    i 
later  on.     This  indicates  ihat  after  a  little  i)iaetice  the  muscles  can  e\ 

eciite  a   uiven   mo\ei t    and   pel  t'oi  m   a    >.dveii   amount    of  \\ork   mm'' 

more  smoothly  than  when  they  are  not   in  traiiiMiir.     Anothci'  intorestiii 
outcome  of  the  invest ijrat ions  has  heeii  to  show  that  work  done  under  ai 

ii"i'i"il  ' iitions  that   tend  to  ]iroduce  any  kind  of  muscular  strain 

done  inctlicii'iitly.  If  has  heeii  found  in  marcliiii}.'  soldiers,  for  exampl 
that  the  slightest  ahrasion  of  the  foot  frreatly  increases  the  ciuml' 
expenditure,  for  the  man,  in  tryinji  to  avoid  the  ])ain  jirodilced  hy  ll 
alirasion.  hriiiirs  into  operation  muscular  groups  that  are  rcall\-  n  ' 
rer|uircd  for  the  elVicient  performain-e  of  the  movement,  lint  are  n^c: 
instead  to  avoid  iiressnre  on  the  sore.  Fatij^iic  also  causes  inciHicic! 
performaMcc  of  work:  that   is  to  sa.v,  the  falijrti.'d  person,  on  attemiitin. 


•  AHIKIV    ll.M.AMi: 


<•■] 


the   sallU'   illlKiinit    of    woik    il.s    lie    ptrfnlllicil    licfiil-c    lii'ciiliiill«    fdtiuuoil. 
nill   till  so  iit   II    lllllcll    L'iciitif  fXpc'iHlitlllc  ((('  tMif|-!.'V. 

Tlicri'  is  n  dimiiiil  \  ;iri,iiioii  in  tiic  ii's|iiiiifi>tv  cxcliiiiiirc,  \\  hirli  is  in 
L'iMcnil  p.-'iiilli'l  with  till'  JMidy  ti'ni|M'ia1urc;  it  rises  duiiin?  the  diiy,  tin- 
Mine  iif  iH'tivity  mid  wmk,  nnd  Talis  dniinj.'  the  niulit,  the  time  of  icst 
and  sleep.  l''ciod  also  alTeets  icspit  jitol  y  exeliaOL'e,  liut  it  will  he  tlime<'- 
i>siiry  to  >ro  into  this  Inither  after  what  has  lieen  said  on  |)au'e  ."(47. 


'        SI 


I 


\..M 


M^^ 


'r' 


<  II.M'TKU  |,\||« 


^Vi! 


A   <  |,|\|(  Al,   MKTIMtl)   luu   KKTKK.MIMN);  TIIK    UKS|'IK.\T<Mn 

KXniAMJi:  IN  MAX 

Mv   H.  <;.   !'i  Ml.  I,   IJ.A..   M.I). 

Principle.  Sinn'  tlir  ilitt'iiiiiiititiiiM  of  ijic  icspiniton  cxcluini.'r  ii 
in.'iii  is  of  Miiiii'  iiii|)<irtiiiii-c  ill  ilit-  stinl\  ot'  crrliiiii  (JisciiM'H  of  tlic  ri'spiin 
tioii,  cirriiliilinii  iilKJ  iiifliilM.liMii.  1111(1  iiU,,  liccjiliM-  ililrctioiis  I'm-  (■iiii\ 
iiij.'  out  till-  iicifssiiiy  iir.Mc.Jiiifs  111 t   irciii-iiillv  ii\ jiilnlilc,  wi-  Imv. 

tllollU'llt    it    MlivJlt    l.f  nf  ;|ssi>ti ,.  f.i   ilwlllllc   llcic   llliff  liil'Cct  i<(||s    for   111. 

Ti.ssnt    illlil    the    DoiiL'hls    iMflliiMJs.      TIli'M'    IIIcIIkmIs    lliivc    lici'ii    t'ouiii!    I. 
(•"iiiiiuiri'   t'.iM.r.ilily   in  iirciiijiry   with   ntliris   in   use  iif    |iics.'iit.t   iilitl  Im 
••iiiiM-  nf  tlirir  ii(ii|it(iliility  {iinj  siinplirity   flicy  nii'  specialty  Miitcil   I'ch 
rlinicnl  unrk. 

I'.y  tlicM'  iiii'tlKHls  the  n.'ru'y  iiicIiiIkiIj  of  the  Imily  is  rjilcilllitc'l  I'ihh, 
(ixytM'ii  i'iinMiiii|iti.in  »y  i-iii'l>iiii  dinxiilc  cxiTctinn  per  niiiiuti>  (iiiiliniM 
.•.iloriiii.-try  )  ipairr  :.4t;i.  tlif  li^finvs  („v  wliirli  aiv  ilrli'iiiiiiii'il  fmin  tli. 
Milunii'  iiiiil  pcrci'iitilc  uaM'iiiis  I'oiiipiisiliini  ni'  tlic  f\piic.|  air. 

Tin- MilijiTt  In-fatiifs  tliroiiirli  \al\<'s  wliicli  aiiti.iiiati.  all  ji.t  ..  i.  n  lli< 
inspiifil  ami  cNpircl  air.  Tlic  fxpiratioiw  fnnii  a  niiiiilii-r  nf  respiration" 
are  e<illeet«'<l  in  a  sjiiromeler  or  liai:.  ami  the  volume  ot'  the  respiriition> 
per  liiililile  is  ileterinilied.  The  ifaseoiis  romposition  ot"  the  expired  air 
is  (Itteiiiiined  liy  nas  analysis,  and  the  oxyyen  eonsimiption  and  enei!;\ 
output  of  the  hody  are  ealeiilated  from  the  data  ohtaiiied. 

Description  and  Use  of  Parts  of  the  Apparatus:  1.  Tm;  Mci  rinii.  i 
AMI  \  Ai.vi.s.  The  inontlipieee  i>  made  of  soft  pure  trimi  riiliher.  and  eon 
sists  of  an  elliptical  riililier  (laii'.'e  liavin;,'  a  hole  in  the  center  2  cm.  in 
flianieter.  to  which  on  one  side  a  short  riihlier  tulie  is  attached.  On  tin 
opposite  side  of  the  hole,  at  riitht  aimlcs  to  the  niMier  tlaiij;e,  are  al 
tached  two  nililicr  inys  The  iiilihei-  llan'_'<'  is  placet  licf.vccn  tlie  tips, 
and  Itie  lin.'sarc  held  li\  ihe  leetii  Tlie  iiiIiIki  tui.e  of  the  inolltlipiecc  i^ 
connected  to  th.'  tiilic  carrxiiiu'  the  \al\i's.  Tlic  nose  must  l>i>  tiirlitl- 
closed  if  mouth  tireatliiiiy;  is  used.     This  is  accimipiished  l>y  a  nose  clip 

\vhi(di  eiinsisfs  of  a   V  sliai)ed  metal  sprinji,  tl nds  of  which  are  in-.. 

vided  with  felt   pads.     A   toothed   la.-het    Is  altaejied   to   the  ends  of  lln 


i'  i 


■Till-  ih.ii.lir  I-  .r|.l..|   |,.r   III. 
■I'.iri'i  !  In        I  .irTirti.     h.-lih:- 


> 11"    .  •    „,,il,ir-   I"   ili^- 

■  f   \\,i-lMin:t..M    k,  |..,ii-.    N. 


!(..    ri|  = 


Ml   lllllh    mil    |.|   II  IIMIMM.     Ill  -lllixll'in      I  M   II  Wi.l      IS      M\\  :i."(."» 

-imiii.'.  ihhI  MTV"-,  t.i  li.iiil  111''  »|iiiim  iiu'liil.v    "•liiiii|ii-l  mi  lltr  imsinU  in 
till'  |ir<i|ii'r  iiiiKitiiiM  I  SIT  I'ju    177' 

Sidiii'    iiidiv  i<liiiils    i'\|ii'i'ii  HIT    meal    il.  lrr>.s    wlu'ii    iiiihIi'    In    lirtiillir 
iliiiiMirh  till'  iiMiiiili.     j-'nr  tlii'M'  it   i«.  Iiot   ti.  iiM'  11   I'iicc  Miii>.k.     I'liruilii 

iiiilriv  al   till'  iiii'M'iit   liiiic  I iisU   K  .ntinly  ^Hti*|■al■tl(|■v       I'riliaps  tln' 

'  .  st   i>  mil'  siilil   liy  Sii'iii',  ( im  iniiii  \  ('n.'   w  hlcli   is  pii'tiiri'il   in  thi'  rut 


I'K      '""         I     ^"-^    .111-.    ".    I.I..     in.!>k.    (,    M..nlli    I'i..., 

\'lii-  liriiii;  |>la I   in   |»»iliiiii   llir   laiT   mask  siimilii   lir  ti'sti'd   I'm-  Iraks. 

Iiii'li  I'iiii   I"'  limir  li\    imttiiii;  •Miap  arminii   llii'  iii'jcs. 

■-'    Till.   \'\i.\is.     Till'   vahi's  i>r  Tissm    ar.'   |)inlia!)i\    tin'   ln'st    I'm'   tin- 

'  ii|Misi-,  I. til   thi',\    ail'  I'NiH'iisiM'  ami  i|iriii-ull   to  i.litaiii.      Wr  liavr  made 

'i-tVi'tly    satistiiftiiry    \ahi's    Irmii    tin'    |iri'|)an'd    i'a>ini:s    um'iI    in    tin' 

I  i.iiuifactiirc  lit'  lii.lii>.'iia  saiisat;''.     TIkm'   ran   lie  ulitaiiicd   piTsiTVfd    !;. 

It,  and  llicy  will  kci-p  indefinitely  mi  iee.     Wlieii  needed  a  slioft  pieee 


■i!*.        ' 


rill-      1.M-k      l:.|.      >„, 

'    \|..ii,ln.„l,     llMi- 


vli  I'-ui  !>    l'\    V  .i.|.,  ■,u  1        llu 


rr^ 


Mid 


:.,')(! 


MKIAMdl.IsM 


is  tliUcil,  «;isliril  iVci.  iVoiil  s.-ilt  liy  ;illn\\  iiiy;  \\:itc|-  rroni  tlic  t;i|i  U,  !i,: 
llii-diiu'li  it.  •■iiiii  sdl'trhi'il  ill  a  uriik  iilv  ccriiii'  sdliitiiiii.  Tlic  iriil  lici-nii!' ■ 
very  suit  an. I  plialili',  ami  .lues  iii>t  ilry  cuiickly.  A  pifcc  cil'  tlir  casji 
iiliniit  III  iMM.  li.iii;  is  tliriMilnl  tlirnii^'li  ii  n-liiss  tulic  ol'  aliiiiit  1.")  iniii.  li..i 
;iii(l  4  to  (I  .•111.  I. HILT,  ('lie  I'll. I  (if  tlic  ciisiiif;  is  lirmiL'lit  anniinl  the  oiitsi.i. 
Ill'  till'  tiiliiiii:  all. I  s.  .-iir.'.l  Iiy  iiicans  .if  a  tlir.'a.!.  Tlic  Ihuit  I'li.l  ..f  tli. 
iiiciiiiiian.'  i.,  iiiiichcd   .ilV  an. I  tin-  casiii"   Is  then   iMit    a   little   iii.nc  tli.i 

hall'  way  aiT.iss  its  ini.lillc,  s.i  tliat   ll |)ciiiiis;-  will    Ii.'  jiiNt   witliiii   li^ 

I'''''  ''"'I  "I'  III"  liilic  w  lu'ii  til.'  i-asiii!.'  is  ilrawii  liacl<  tlifimjili  it.  'I'i 
l.M.sc  end  i.t'  the  casiiiu'  is  sliolitl\  twist. ■.!  an  es.s.'iitial  proce.jiir.'  an  ■ 
is  til. 11  sci'tiii'.l  liy  a  tlir.'a.l  mi  tlie  i.iitei-  si.le  ,.|'  llic  tulfe  If  |ir.i|.,.il. 
ina.l.',  til.'  val\.>  will  w.nk  iV.M'ly  without  \  iliratimi,  and  f  lie  ..p.'iiiiM.^  l.- 
siifdeieiitly  lai!je  to  aIN.w  a  jf.iM.l  cnricnt  .it'  air  to  pas.s.  It  sli.niKI  .i,! 
la|ise  instantly  an. I  lu'  air  tii;lit  when  tli.'  current  of  .air  is  reversed.  Tl  ■ 
hack  lash.  ,,r  lau'  •<(  .•losiii.',  of  these  valves  is  extr.'iiiely  small,  an  ' 
th.-y  will  .i|'eii  or  el.'se  with  a  pr.'ssiir.'  of  air  n.it  exeeedim,'  tli.'  p|-.'sMr. 


I'lk:.    i:^      I':. 


I.'i  U.lti'I  >      \.ilv<  -. 


i- 


.   1 


li 


i^  ^1 


ehan-.s  ill   inirnial    respiration.      Wli.Mi    not    in    lis.',   tli.'   valves  should    ! 
kept   in  ;:l\,'.'riiie  wat.M-  .hi   i.-e.     \alves  pr.-pared  in  this  way  have  h,  . 
in  lis.'  a  iiioiitli  with. .lit  l.iss  ..f  ..fli.-icn.-y.     They  are,  liow.'vr.  nia.j.'  wi- 
s..  -r.at  .as,,  that  ii.  w    \alv.'s  aiv  pn.vi.l.Ml  f.,r  ea.-h  siili.i.-.'f,  an. I  they  -. 
tlii-r.'f.,re  .•>! iaily  a.lapte.j  to  war.l  w.n-k   l  l-'i^'.    l7.Si. 

''"'"■    ^.'ilvi-s    are    ins..rte.l    in    r.'vei-,'    .inhr    into    :■    support'im'    iii.  t. 
T  pi. ■.•,.,  an. I   tl..'   .1. lints   ma.!.'   aii'  li-hl    liy    tap.'.     The   stem   ,if  the  T 
'■'"""■'•''■''   "I'll   III''   IN. iiithpi. ■,','.     Thi-oiiudi   a    rnlilicr  liilie  of  ahont   .'l 
iin-h  hoi-.',  th.'  .xpire,!  air  is  .-olhct.'.!  in  the  spir ter,  or  |)oii<rlas  I'.a. 

■•■  Till  Ti>MiT  Snii.-.iMi  hi;  is  pi.-i,iiv,j  i,i  i-'i;,  17.).  \v,,  |ia\,.  f,,,,,,,}  ti 
'""  ll''-''  -''■'"■  I"  !"■  v.'iv  s,.r\i.-,'alile  in  ih,.  ..lini...  This  iiistriim.'nt 
in"iinl.'.l  on  a  platform  la\in._-  riilil  .'r  w  h.-.'U.  an.!  .'an  I.e  iiiov.'.l  ah.e 
the  war.js  with  .as...  Th.'  h.'ll  .if  th.'  spir.unel.'r  is  nia.h'  .if  aliiminn' 
ami  is  snsp,.n.|..|  in  a  wat.iliath  li.'lween  th.'  doiililc  walls  of  a  lioli.. 
evlin.l.'i-  mail.'  of  uahani/.  d  ii-n.  Th.'  liei-lil  „f  ||ie  |„'||  i.s  7i.>  ,., 
"'"'     ''i''     'liani.'ti'r     Vl     .'m.        An     ..pi'iiin-r     at     th.'     Ii.ittoiri     of     :' 

e.viin.l.'r  .•.mne.-ts   thi'.ei.ih   a    tli! -way   stop>-o,-k    v.  it),    >  !,„   !-.i|,)HM 

l'''i''i'il^'    l'i'"iii    1 1 \piralnr,\     \al\.'    ..f    th..    moiithpi.'.-.'    is.',,    l-'i^.       TT 


MITlliili    KOU    liKTIinilMMI    lilsnifATuiiV    i;\rll.\.\(,i;    IN     MAN  "i.")" 

Tlic  licll  is  I'lHiiitcipdisi'd  liy  iiic.-iiis  lit'  ii   wciulit.      In  tlic  (iriu'iiiiil   Tissn) 
iiiriiiiiclci-  iiii   initniiijiiic   jiil.iusliiicnt    pcrinittcil    \\:i\vy   in    .•iiiiuuiit    (■(|ii;il 

'..  tilt'  \\;iliT  ilispli 1   liy  Ihr  IhOI   til  tliiw   t'luiii  llii'  s|iii'iiMii'ti'i'  cylimliT 

::ilii    a    ciimitci-ixiisf    i-yjiinlir    ;is    tlu'    licll    asi  i.niicil    mit    of    tin'    water. 


Ill'  lii'll,  lii'iiiir  licavicr  out  nf  watn-  tliaii  w  hi'ii  it  is  iiiiiiU'i-scil,  is  accufil- 
■aly  ciiiiiitiTpiiiscil  ill  any  imsitiim,  altliniiu'ii  t 'arpi-ntcr  lias  slmwn  tliat 
is  pcfiiii'iiii'iit  is  iinnci'cssary.  An  upciiinji  in  flu'  top  ui'  tin"  spiniiiicti'r 
iiiiits  tlic  iiiMTliiin  III"  a  nililicr  stuppcr.  tlirninj-li  wliii-Ii  arc  passcil  a 
i-niiiuiu'ti'i',  a  water  iiiaiiniiit'ter,  and  a  stupcoek  with  tube  I'lir  draw  inir 


41 


i    ir 
1    ■> 


iill 


il 


I 


1; 


J    I 


1^' 

t  ■  1 


1;^ 


'"11  1 


11 

4  ^ 


y 


.(.iS 


Ml  r Mil II  IS \i 


llii-  Niuiiplr  nl'  air.     A  sralf  cm   i|,r  sidr  i,\'  till'  iiistiiiinciit   i;iv.'>  llic  \,,| 
uiiK'  of  tlio  air. 

Diiiiiiji  an  iiliscrvatidii  the  siilijccf   siu  in  a   rrc-liniim  pusiticiii  up  i;., 
npdii   a   cipucli.      WliiMi   tlic   hell   (if  tile  spiidiiii'ti'i-   is   placed   at    /cin.  tl  ,■ 

iiiiinllipicrc  adjnsicd  in  llic  nlli    and   tin'  ni>M.  i-lainpi'd.  rcspirat  idn   is 

started,    llie    e\piial  ioiis    liein-;'    passed    llinini;li    tin'    slnpi k,    \\lii,-|i    i, 

sii   turned    as    tn   allcjw     tlieiii    to    pass    t.i    the    imtside    air.      After   a    n-^ 
lilinutes  the  stupe, ii-k    is  turned   s(i   that    llie   expiratinns   are   passed    ihi,, 


'■'«■. 1^"        n>,      llontl.!-    iMf    in.n,..,l     l,„     ,l,t,,„„,nni:    il„      ,,  ,|.,r..lc.i  v     ,v.lui„K,..       Th,      „-. 
■Ufiil    ..t    iTi.Mill';^.,,.    v,,K,~,    ^in.l    ,,.iii„,t„;„    I, ,1k..    .|,,.„„     h.r,.    1,,,.    l„,-„     f,„,,„|    ,,,    |„.    ,„,,r, 
M-iniiU    llKin    that    i .  ,  ..ini.i,  r.|,  .1    l.i     l).,ni;l.,, 

the  s])ini|iietei'   for  a   delinile   hiiM|li   i,f  tiiin-.      At    tlie   end   i>(  the   peii     ' 

'li' '■!<  is  auain  tuined,  ami  after  the  liar.nnetrie  |u-essiire,  teinperatn  • 

ii'i'l   Milnn f  the  air   havc^    Keen    noted,    iln'   ei.iiipii>ithin    of   the   air     - 

determined  in  the  ilaldane  i;as  analysis  apparatus, 

4.   Tin:  I)nr.;i,As  \'>\i;.      The  Dnnulas  liau  is  made  nf  luld.er-lined  ,-1..' 

iinil  is  eapahle  <.f  hi.l.liiii;    fr .Vi  t<i   lilii  lit,. is.      It   is  os|)eei;ill\    us.  ' 

for  investiiraliuiis  diirinu-  .•\er.-ise.  sinee  it   is  litleil  with  strnps  so  i! 
the  liJiiT  can  he  fastened  to  the  shonl.jeis  i  I'iu.  1,^(1..     It  is  then  eenniM-i    i 
uitii    tin'    valves,    tlir    mnUt  lipieec    of    which    is    pl.iced    ipelwcen    the    Ir 


MITIKIF)    Fill!    Ill   TlliMIMSd    lilSI'IK  \  Tl  (If  V    IN 


III  VMil;    l\     MA\ 


IfcNpiratiiiiis   iirc   i-dinnn'Mr.-il    wiili    tli..   ilncc-wiiy    vnlvf    iiinii-cl 


;illo\v  the  cNpinif iiuis  to  |i.    s  ilirt'ctly   oiitsidc.     At't 
liliriiiin  is  cstiililislicil,  tlic  ilnrc-w 


Ml      lis      til 

«'!■   icspiiiitiii'v  (>(|ui- 


!iy    Vfllvi'    I-.    tUIIKM 


I  ilii 


liii-v  ])(>i'io(l  Ml  tliat   ill- 


niifj;  III!  msi)irii- 


'"•  ^ii< <'<liiitr  fNpiiiitioiis  iiijiv   i)iiss  into  tlic  lias.'. 

Tlic  time  iiMiiiiicil  to  till  tlic  luiir  iMniilnit.ilily  is  iIctiTiiiiiicd  w  itli  ;,  slnp- 

\\iitcli.     The  nir  vliii-li  li.-is  I ii  (•(illfi-icij   in  the  liiiir  iliiiiiijr  the  pcriini 

is  tlioi-duclily  inixiMJ  mill   piissid  tin-diiuli   m   niflcr.  tiic  tnnpciiitiiic  aiiij 
lijininiftiii'   pri-ssuii'  .ir.'   imtoil,  :iiiil   a   sample  aiialy/i'd   in   tl,,.    Hiililaiic 


'Jiisappaialiis.     The  Iia-r  slnnild  Im  ciniitiod  coinplotcly  liy  lolliuff  it  tip 
"lien  iiparl.\-  pini)ty. 

■')   The  Haldane   Oas-analysis  Apparatus.      1'kim  ii-i.i;.— Tlic   llaldain> 
''i.tliixl  (if  analysis  of  cxpiicd  air  Is  simplr  nnd  easily  Icani.'d.     TIip  ap- 

'larattis   ( Fjjr.    l,s| nsists  „(  a    "ias   Iniioit.",   a   ciiiitinl   liuicttc   of  tho 

ainc  size  (li.ith  surroniidcd  with  a  water  Jaclieti.  and  Indlis  .•ontaininsr 
iilute  caiiNtic  jMitash  m-  soda  soliiti.in  for  the  Mosoijition  of  the  i-ar'.  ti  di- 
Nidc   and   an   alkaline   pyi-oi:allate   solution    for   the    alisori)tioii    of   tho 


1^ 


m 


^1  J 


ii^ 


^  f'. 


if*!':' 


'J;'  ■ 


it     I 


it ! 


fi  If  I 


^J!  i 


:i*>() 


Ml.T.MidT.ISM 


oxyiicii,      Tlic   u'Ms   Imictfi'   is   (•(iniKctcd   witli    tlic   luilbs   liy   ji   t\M>-\ 
st(i|iciicU,  \\liicli  iillciws  ,-1  sMiripli'  lit'  ".'.-IS  to  p.iss  iiitii  cither  Inili).     A  . 

ti'cil  tiiiii'  I  III)  is  put  iriti)  iMiiini'i'tiiiri  witli  tlir  Imn'ttc  iliroiijrli  ii  inini 

Illlic,   wliii-ii   is  ciiiiiici-trii   uitii   till'  ;ilk;ili   Imlii,  ;im<1  ci'i  ln"  nijiilc  to  cni 
piiisiitc  fill-  iiiiy  i-li;iii'_'rs  in  l('iii|'fi;itui-('  tii.it  iri;iv  niTUf  diifiiii.'  tlic  cmii   • 
i<f  tlic  .■iii;ii\sis.      I-"iir  ;iii   aiiMlysis  llic   i^.-s   is   ti-MiislVrrcil    ti>   tlic  Imiic!- 
I'nini   the  saiiipliiiii'  tillie.  satiiratcil  \\itli  watci'  \ap(ir  ovei'  iiicrcin>-.  ai,  i 
then  iiieasiiicd.  aftiT  which  it   is  t I'aiist'circd  into  tiic  caustic  sdhitiun  t- 
free   it    frniii  CO.,  and    rctiinied   tn  the  Imrctte   tn  dclerniiiie  the  luss  >■ 
Milmiic  dii.'  til  ('(I     alisiii  ptiiiM.      It    is  then  tiaiisl'crrcd  iiit.i  the  alkali'- 
pyruu'allatc  sdliitinii.  which   frees  it   fioiii  oxysicii.  after  wliieli  it  is  auM; 
lu'.ini;ht   liack   tn  lli,'   liiii-cttc  In  detciniine  the   hiss  in   vuiiinic  due  tn   ih 
alis(ir|i! inii   iif  the  ii\\itcii. 

Tin:    AervKxi'is.      The    detail    nf   the    llaldaiie    apparatus    is   slinwii    i" 
the  accMiiipanyiut;  cut.     The  nicasuiin^  lunette  i  / 1  holds  i2|  c.e.     Tic  Ipiili. 
Is  of   I.'i  c.e    eapaiit.v,  and  the  j,Maduatcd  stem,  which  is  alioiil   4  iiiiii.   ii 
hole  ami   till  em    ill   Iciiirlli,  is  iiradualcd  to  (l.dl   c.e     from    1.")  c.e.   to  _') 
!■.•■.      The   slope.  (•!;    ;;1    the  to))   of  the   luirette    is   ihuilile-hored,   so   that    i" 
one  position  ail'  can  lie  drawn  in  from  a  uas  sampler  i.'i   and  in  aiiothr 
^'■i>'    iiilo    lie-  alisoriition   hiilhs  (Vi,     The   lower   jiart    of  the   luirette  i\ 
teiiils   throiiuh    the   nililier  cork   at    the   liottoni   of  the   water  jacket    i  / 
A   piece  of  rnlili.u'  tuliiim  is  altaelied  to  the  hottoin  of  the  luirclte  ani 
is  passed  thiiPiiLih  a  nieial  tiihe.  furnished  on  its  inside  with  a  metal  d's 
uhieh   presses  au'ainst   the  ruhlicr  tuliiiii:.  the  pressure  licinjr  controlled   I 
means  nf  a  line  ailjusiinir  screw  >ii>.     I'.elow  this  a  ylass  stopcock  (;  >  ei.i 
nects  u  ith  i-iilil.er  tiiliin!,'  to  the  iiici'eury  Icvcliii';  liulh  (.i  i.     The  ahsorplin- 
luilh  for  ('<»..  eoiitainiii'j:  "JO  jier  cent   Nadll  or  K<»|[  (.<>;.  is  ])ut   in  coi 
iiei-tinii  wiili  th,.  liiiretle  liy  suitalily  lurniii!,'  stopcocks  (.,'  and  >>'.•     Ti' 

• 'rol    liiiii'ilc    i/ni    is   also   iu    connection    with    this   hull)    throu'.di    ti- 

manoiiieter  tulic  ill 'J      .\i\\    variation  in  temperature  whiidi  may  ii 

diirinu   the  analysis   will   cause   the  level   of  the  alkaline  solution    in   tl: 
manometer  to  clianjr''. 

When  final  readings  of  the  sliriiikai;c'  of  volume  are  made,  the  level  ■ 
the  i-ailslie  s.jitlion  is  returned  to  the  hvel  of  thai  in  the  manoiiiel. 
hy  so  doinj.'  aii>  error  due  to  temperature  chanijes  is  avoided,  siic 
i-haiiL'e  ill  tc'iiperaturc  must  he  equal   in  the  twd  hurettes. 

The  ahsorptioii  Imlli  for  oxytreii  ( /,M   is  filled  with  a  solution  made  I' 
dissoKiiivr   in  yraiiis  of  |iyo!.'allie  acid   in    100  c.e.  of  a   nearly  saturat. 
I\(HI  solution.     The  specific  frravity  of  the  K(»II  should  he  1. ;")."),  which 
ohtaiiied  approximately  hy  dissolvin;,'  the  sticks    (pure  hy   alcohon    in  a 

'Til.-    ^ti.i......  L     (  ^  1     ,.     .]   ...I.t.     l.,.r..,!.    -.,    e.;;;     t),.      ...I,.-    I..  .,1;..^    fr.!::    ;!:.      *..:;:  .  t: .  .::     !..      ! !.. 

Cwnit.  '  tpiM     uilh     «  Itlui      ''     MI      1  J 

trills  liibi-  .ils.i  h.iK  a  thicfway  sl..ipc<>ck  I  niK  so  ihat   it  may  he  i.|,i-ncil  In  Hit-  cmtsi.le. 


Hi        'l! 


MITIilPl)    niU    lUTl  inriNINi:    HISI'lliATDUY     l.X(IIAN(;i:    IN-     MAN-  Tlfil 

('i|iial  \vi'i>_'lil  iif  uiitcr.  Tln'  iiiarU  ( i:,)  on  tlic  stem  of  tlic  I>ull»  iiuli- 
citcs  tlic  lc\(l  ;it  wliich  tlic  sdliiliiiHs  sliniild  staiid,  Kiioii<rli  pyroiriilliito 
^Hliiiioii  is  intiip(lui-.'(l  tlndiiij:li  liilio  /."  to  fill  Inillis  /.'  and  //'  two-fliirds 
I'lill.  Tlicii  pvioirallatc  solution  is  |)o\ii('d  into  tiilic  III  until  tlic  difTcr- 
1  lice  ill  lc\c'l  ol'  tlif  lliiids  is  sufficient  to  jiidiliicc  cnoii>;li  ])r('ssure  to 
laisc  the  lc\('l  of  tlic  pyrotrallatc  solution  in  /,'  to  tlic  level  /.'  on  tliP 
siciii.  Stopcock  s  must  he  open  duiiiiir  tliis  iMdecdiiie.  It  may  lie  neeos- 
saiy  to  add  or  tale  a\\a.\  a  little  p.\  roLrallatc  solution  tliroufrli  /■;  to  at- 
tain tlic  aliovc  Ie\('l. 

('ai-e  mu>t  lie  taken  to  allow  for  complete  alisor])tion  of  oxyu;oil  from 
ilic  air  tliat  is  entrapped  hciwccn  //  and  /f,'  liefore  an  anal.\sis  is  made: 
otiiciw  isc  cliainjes  w  ill  lie  )iri.iluced  in  tlic  Icxcl  of  tlie  iiyrotrallafe  solu- 
liiiii.  'riie  air  in  tin"  capillary  tiiliini.'-  i-oniicetinu'  the  liiireltes  with  the 
al'sorptiiiii  liiillis  must  also  he  freed  of  ('(I,  and  <»..  This  can  he  aci-oni- 
plished  h\-  niakins.'  a  ilumiiiy  anal\sis  of  atmospheric  aii-  hcftirc  the  real 
analysis,  (in  at  care  must  lie  taken  to  have  atmospheric  ]>ressiire  in  all 
ilie  tuhes  at  the  start  of  the  aiialxsis.  This  is  a<'eomi)lished  liy  opciiiu),; 
the  st<ipcoek  in  the  hniette  first  to  atmospheric  air  and  then  to  the  ali- 
siirption  hnlhs.  until  no  further  (diaiiire  in  the  level  of  tht>  llui<1s  in  the 
stilus  of  the  ahsorptii.n  hulhs  i ceiirs.  This  level  is  then  marked  and 
used  as  the  standard.     A  small  amount  of  \xatci-  in  the  hiiretfe  over  the 

rcury  assiins  saturation  of  the  air  with  water  vapor.     Time  for  draiii- 

ai;e  iiillsf    he  allow  eil   hel'o|-e   liiakiiie-  readiiiLrs, 

.\  very  ser\  icahle  mtiiiiiliiKj  liilii  for  the  transfer  of  air  can  he  m.nde 
iVciiii  a  :'><!  c.e.  eronnd'_dass  syrinire,  to  whiidi  is  attached  a  two-way 
•-topcock.  A  cut  of  this  is  shown  in  l-'i;;.  1^1.  The  ilcad  space  in  these 
^yriiiLres  is  washed  out  hy  uorkimr  the  jiisfon  haidx  and  forth  several 
tiiiii's.  A  thill  coatiii<_'  of  vaseline  iircvcnts  leakaue  of  the  <ras.  \Vc  have 
ioiiiid  that  these  saiiipliii','  tuhes  will  retain  a  sample  of  expired  air  with- 
out eliaiii:e  up  to  eitilit  hours. 

.AFami'i  i.ATroN  (IK  Ai'i'AKA'n  s.  The  saiii)ilinn  syriiiy;e  i  .v;'i  is  attached 
I"  opeiiinir  ,'  (if  the  hiir.'tte.  and  its  sinpei.ck  i  /,"  >  opeiieil  to  a!mos|ilierie  air. 
The  Ii-mI  1,1'  the  iiierelir.v  is  raisi'il  to  the  level  of  the  stopcock  of  the  syringe 
iiihI  is  then  tiiriird  s'l  that  s\  riiiire  and  hiirette  are  in  coiiiiiiunieation.  The 
"iiUi  of  mercury  is  lowered  so  that  the  mercury  falls  in  the  hurcttc.  Th's 
draws  the  piston  of  the  syriiiMc  with  it,  and  fills  the  hurette  with  air 
lioiii  the  s.\  riii^'c.  it  is  advisahlc  to  jint  a  little  jiositivo  ]iressui'P  ok  tlie 
pistnii  of  the  syrinire  in  the  iiianemcr  to  prevent  ])ossihle  Icakajro.  When 
all  of  the  air  is  in  the  hiiretle  a  slielit  |)ositi\e  i)ressui'e  is  iircMltieed  in 
n\e  hiirctte  hy  '.M-otly  [o-.  ss!e,n;  nn  the  piston,  and  iiiinie,!iately  there- 
after the  stop k   on   the  syrinee   ( /,"  i    is  airaiti    turned   to  tlie  oi'ininal 

p'isiti(jn.     This  allows  the  pressure  of  air  in  Iln'  hurette  to  com.'  to  that 


I 


Iff 


V 


: 


\'i^\\\i 


i  '■ 


!-=;  , .  i 


4 


562  MKTAImt.ISM 

I't"  till'  Mt s|ili(ic      Tli(>  Iii'lirlil  (if  till'  iiicicury  is  now  adjusted  to  a  ■■■■i. 

vi'iiifiit   liciL'Iit   ill   the  liiircttc  \i\    .•icsinir  ,.|ick  7'  niid  tiiriiiii!.'  the  mill.' - 

screw  '.'.     Till'  k   l>  is  ik.w  nia.I,'  to  (•(iiiiiiinnicatc  wilii  tlio  al)soi|itii. 

Imllis.      II"  tlif  air  ill   tlii'  Imrcltc  is  at   a1iii(>s|ilifrii'   prcssun".    no  cliairj 
"'"  ' Ill'  ill  tlu'  l.xcl  (,f  111,.  iliii.K      Tlic  niidiii!.'  is  tli.-n  tal<on  on  ili. 

Illll'fllC 

Till'  lic\l  >t«ii  ill  tlir  aiialv^is  i-.ii.vivis  ill  tuiiiinir  st.i|ii',H-l<  s  to  coHi 
imiiiicat-'  uitli  llic  .■.iiisti,'  voda  -..>liiii,,ii  in  I, nil,  !>,  „iid  tlio  IrvcliiiLr  iiil„ 
('^  is  raised,  r(.iviii!.'  iii.T.Mii-.v  iiiln  tlir  Imiii'IIc  and  tin-  aii'  into  liiilli  • 
Till'  m:\:  is  iiasM'd  liai-k  and  forlli  s,'\iral  linn's  until  absorption  is  coii! 
|>li't.'.  as  I'an  lif  drli'rtiiiiii'd  l,y  lln'  fact  that  tlii'  livi-l  of  tli,'  mi'miry  ii 
tlio  liiirctti'  ri'iiiaiiis  ronstant  wlicn  tin'  llnid  in  tlii'  Imlli  is  rctiiriu'd  f.. 
its  oriirinal  Ii'mI  /.;  ,,i,  ih,.  st,  ni  In  lliis  adjiistnn'iit  it  is  .•onvi'iiini' 
to  mnko  till'  -less  li'Vi'li'M.  l,y  III,'  iiii'ii-niy  Imlli  and  the  line  li'vi'liim  '■ 
I'losiiii.'  r  and  liirninu-  i.  until  i|i,.  Unid  in  .'/  is  at  llic  oriiriiial  Iii'IliIii 
Till'  i-i'adiii-  on  111.'  Iniiitt.'  indii-ati's  tin'  Inss  in  volunn-  diii'  to  tlic  I'd 
alisorlii'd. 

'•"' x>Ui'ii   is  iiiiioM'd  li\    ;i   similar  iiniiTiliiri'.  tlic  iras  lu'liiir  i>ass,..! 

into  till'  all;alini'  |i\  roirallali'  solution  liy  tnriiiii'.'  mrk  s  to  i-oiniiiiinii-ai. 
\vitli  Iiiill,  l.\  Till'  alisiirptimi  oi'  on,\  u.'ii  is  slowfr  llian  for  CO..  an! 
nnir.'  I'ar,'  niiist  !ir  taUi'ii  lo  uil  i-i,nii.li.t.'  alisorption.  Tlic  air  in  tl  . 
tiiliin-  liitu,'.  |.  till.  il;,i,|  in  /-  aiid  sto|..-,„.U  s  nmsi  |„,  waslicd  out  sr\ 
cral  tinii's  in  order  lo  m|.|  tl,,'  owucii  wliirh  is  left  in  it  after  tlie  alism-i. 
lion  of  tlie  CO  .  ■VVIien  iMs  is  eoiiiplete,  tlie  tinal  I'eadillU'  on  the  lilirel'. 
is  made  and  the  loss  in  \olnnie  from  the  seeond  readiiii,'  represents  th' 
o\yi,'en. 

THE  CALCULATIONS 

Thr  iiiJriiliilloii  tif  til,    i„  rr,  iilili    ,„,i,/«,xi7/„»  „/  Ih,    air  inul  iif  tin    r, 
xi>!ral(,n)  tpmll,,,!   is  represented    In   the   followin-r  example  of  an   aetii.r 
analysis: 

fTlie  tcniperntiirc  and  liaioniel  i  ie  pressnre  as  taken  al  tlie  time  of  Ih. 
cNpcriinent  were  20'  C.  and  747  mm.  ll<;.'i 

1st   re.'eliiij;  of  liiirolti'    20.00 

L'n.l   ii-.'i.liii',;   nf   l.iui'it,'  :,ri,i    i,I,>.m  |.i  ii.n    .1'   CO VX[in 

('(^,    :il.s..,l.,.,l     _    T^J 

O.SO-:- I'D  — -l.il  |„.r  ,M'iit  CO    in  eN|,ire,l  tiir. 
(),  iiiiiilii'ii'i 

2nil    rcailiin;    of    lniiilli' 10.20 

.IriJ   rrri.Iiii;:  nf  lnuctlc  Mi'lir  m1.<..i  |.tl..ii   ,,1    o '    l-,no 

O.  alisnrlii'il    3  -jf) 

'■'.'.'■('    :     I'M--  K'l.riii  |.,'r  I'iMil  of  (I    iTi  i'\;iiii'i1  air. 


Mi'i'llciii   I'lti;   HI  11  i;\iiM\i:    iM^ni;  \  rniM    i\iii\si,i     i\    \i  w 


III  If  iiuimilii'n  I'l'  A'.',' 

O^   in    iilnh'splicii.'    ;,ii  Sn.OK/J, 

<).     CO,  ill  cxi.iir,!  :iii      ir.r.m    li     --.  2(t..",iK;, 


■2'K:'\ 


\    II:    ;,ti 


"-I'lirl  !■'  Mil 


111    i'\|.|li'' 


Since  till'  iiiliiiiii'ii  is  lint  I'liiihuiil  ill  Miliiiiii',  till'  1;in1  limii'f  sliows  tli;it 
Hire  ci\yiic'ii   must   li:i-.r  lnin   l.-iki'ii   in  iliiiiiif;  inspiviilinn  lliiiil  O.     ('(I, 


,i-i  Ih'cii   liixcii  liai'li   ill   !■■  "iral  iiii 


'I'liis  iili\  iuiislv   iMilst    lie   iMJii'li    ilili 


I'liiiiit   ill  tlio  ('alriiliiliiins.     Till'  aiiiiiiiiit  111'  <^  ai'linill>    iiKpii-i'd   I'm' 


|iMI  c.f.  111'  air  I 


'Xl)iri'(l  is  I'l 


iiliiil  as  tuljnws: 


.'".;•  I 


I   ','       (),      ill      .'itllHlSlil,! 


;i.li(i   c^c   N,  ill  •■iliiMisplioiii-  .-lir; 


lit   liM-tMr 

iM    llMXV   I 


i\t'  limi 


■'.'.."i    .  •  ,     \    liiiili.i    I'll    tin 
I'll  -nit   ill  r\]  iinl  nil   t-i 


.■1.|'7,   III.'   \..li; 


II. I'll.)    

..f  (I    ulii.li 


ml    f..l..\    l.l.^-rlit.l 


.'I  1 17     iii.rifi_     I., 


0     :i.|ll:lll\    lil-oil.r.l. 


II     n-i'i 


1..I 


.■tr.j. 


I. ".7 


.  Ill'  ii'-|'ii;il"i>  •!  :iiti.'iit.  11!   iiiii"  1.1  I '( •   .viri'l.-.M:.  <>    :il.s..il,c,l. 


Tiilcl    h'ds    K.!ili,iiiii(.      Till'    vulliiiic    of    ail-    r\|iilri|     ill     l."i    millllti's    illtii 
■hi'  'J'issiit  spiniiiii'ti'i'  was   rmiiiil   In  lir   lt"l  liti'i's  iiicaMiri'il  at   !-'••     ('.  ami 
717  iiilii     Il^r    I  lirass  si-ali'  liarniin'ti'r  * .      'I'liis   vnliiiiu'  nl'  -.'as  must    lie  nir 
iirtcil  sii  as  to  L'ivi'  till-  \iiliiim'  111'  dry  air  at  II     and  TtiH  liiiii.  llg.     To  dn 
'his  two  tilings  must  lie  laUni  iiitu  a unit.     •  1     Since  tlic  expired  air  is 

•  iturated  willi  water,  the  ])re>sure  due  tn  water  vapor  must  lie  subtracted 
:i-<iiii  the  ()liser\('il  liaviniiel  rie  piesNiin-  In  iiliiain  the  true  itressure.  The 
\apor  tension   of  water   I'or   \aiious   temperatures   is   jjivcu   in   Talile   II 

■n  pa^e  ."id-l,.  1 -J  I  Tile  liarniiieier  tiilie  leiiM'i||,.|is  iif  contraets  with  heat 
■  ir    eiild,    and    therel'me    the    liarniiiet rie    icadiims    iii'iist    lie    corrected. 

riie  cori'cetions  I'lir  unlinaiy  liarnmeti'ie  readinu's  are   ''1  und   in  Tahlo   III. 

M'J-i'     ."id.'l.        The     li'.;lire     eiiri  rspnlldili'J-     to     tile     I  elll  per,    ,  U  fes     is     slllitraetl'd 

'iiiiii  the  Iiariiiui  1  lie  readiiiy  in  order  to  nlitaiii  enrreet   lianimetrie  iM'os- 

■'ire. 

In  the  aliove  e\])erimeiit.  the  correetioii  Tor  the  harometer  is  '.2.41  itliii. 
^1  e  Tahle  III.  pa'^'c  ."Hi.'ii.  and  that  I'm-  sapnr  teiisiuu  at  2(1  < '.  is  17.4 
Mv  Tahle  11.  patre  .'.(14  >. 

AiliKil  Iliii-'Diii  li-i,  /'(■,  vMU'  747  i  17..')  -J.:!!!)  7"27.'J1  nun.  The 
iietlicient  ol'  exiiausion  of  L'ases  is  taken  as  (I.OOlitif!.")  1  or  1  27:1;  thei'cfoi'c 
he  volume  of  0     eipials  the  \  olume  at    1     divided  Iiy    I      0.00:'>(i(ir)  t;  and 

lelU'O 


■  riiis   t-   til"  c.pii-t.iiH    Ci   i'l  rt<  r.t.inc   'n   .I'r 

•  l"l"-  i-;iUul.,li,.»    ,;;•!    1i.-   -rii|, I    \.s    M-iiiLi    ,,ii     I'.Ii  n  i.il.-.l   l,ii!,'    i  ]•. 

ii'ironditig  to  tilt,    v.lii'iiis  ftrn  iiKki  •;  (>i   N    in   tin-  iviiir.l  .nir. 


.11    uiiim;   til,-   II. 


I 


|i 


!h: 


'^ 


:H=      I 


i:- 
Pi 


I  •;  i 


:|:l'i 


Ills  n-' 

li  j  i 


r>(;4 


Vn 


\-  .  i;:.-!      ^    V 

-7:!  '  t      I  ■  (i.dd.'Kiii:,  I 


MITMlOI.ISAr 

,  Hh.ri   \„        \-,,|ii,iir  111  (I    :iii.|  \-        V„l, ,,, 


Till'    l"lu I'   «:is    ImmII-     illMTM'lv    !,.<    111,.    |,|,.,,,|U1V    \'( 

l"  I'lvs^urr:  i.r  Welkin^  Ldli  r..iii.,|i,  i,«  |,.-|.|||,.r, 


7(H! 


ulini'  V  ~  Milii 


V.>—  .. 


Vl- 


VP 

lid' 1 1"  o.o(i:;iT(;rrTi 


iiiii  ■  (i't::  •  I 

Tliis   r..iiiiill,i   ;i|)|)li.',|   t(i  tlic  l)rfs.'iit    pinlilciil   lC!l<ls: 

1-,,  liiii     7:.':.i; 

'"'-"..,  -  ^'.<::  bills, 

.iiii  (,1  .  (i.iMi:;i;t;j  ■  -^Oj 

Tlir  l.-illcr  cjll.Mll.ltioM  .-.•111  l„  (■nilsMl..|-iil.|y  silliplilicd  ],y  nsilij:  st.in.l:ii. 
tallies  wliirh  uivf  (•(instjiiits  lor  (•ornvtiiins  of  <.ms  Vdliiiiifs.  Tlicsr  ,ii 
c.-iNily  .liitaiiialiji-  ami  arc  irivcii  in  jiart   in  Talil(>  IV. 

Accinliii-   to   tlii's,.   tallies    for   L'l)     c.   ,,11,1    Tl'T.L'l    nun.    H-    ll.l'..    il, 
factor  is  (l.s!)l-_>4;  tlierefore: 


l'>:'IJI  •   im,        yi.io,  ,1,,,,^    ,,  ,.   ,^|^^,  .|.|,  ^^^^^^    ji^^ 

">:'IJ1  ■  I. .-,7    -  III.7  lit..,^  „|-  ().   1,1  1.-,  ,„i„ i.Tjs  (..  ,„.,.  |,,„|,, 

Thr  Cilori,     Vnhi,    Cilriihit, ,1  from    lln    (!,i,    Krrl,„i„i<.-\W    refer 

to  Tal.le  V  i.'ivinir  the  heat   vail f  I    liter  of  (»,  at    various  'respii  aln,  - 

'Piotients.  it   is  founil  that  at  a  UXl  of  (I.,s7.  4.,ss,S  eaiories  are  ex,ieii,|,.,l . 
lti.2S   liters   <if  <1.    is    therefore   eiiuivaleiit    to    IS. 4      4..'^S,S         7!»   eaiories 

The  results  must  he  ealeuiate.l  for  surface  area  as  well  as  ImmIv  weiulit 
Siipiiose  the  snhjeet  wei^rl,,.,!  s",  kjr.  ,.|||,1  „as  170  em.  in  hiN-ht;  l,v  leh' 

eiiee   to  the  chart    for  i leteriiiiniii-  th.^  siirf, iirea   of  man    I  |.ai,'e   .Mn 

tllis  wouhl   lie  fouml  to  lie   \'M\  square  meters.      The  caloric   expemliim. 

I'cr  s.|uare  meter  in  the  aliove  case  is  therefore     /".     .40.:!  caloiies 

l.!l(i 
Tmuk  I 
Tin:   I'M.iKMA,;,.  ok  (Ixsi.in   whi,  1,   ,s   Koi  ivm.knt  to  thi:  Nni;,.;iN   Kimn,,  .^   •,■,. 

K.\l'li:iTi    .\||; 

T"  ';'•'•■''"  ','"■  ''i""-''"   "I   II \|'ii..|  Mil,  .-1.1,1  111,,  |„.n.,.,in,.,.  „|-  CO    ,„„|  ,,    ,■„„„ 

i'lo!  hiure:"       "  """  '"""   '""•     '''"■  '^''''"  "'■"^  "'"  !"' '•'"•^'«''  '' '"--I '1-^  '  ' 


'^^^     .;'■'.  ;'"  ■'"  ■"■"  ""■'  ■"■-  "''■■■  "i'-t        -'■'■■>       7!M!       7i.7       7" 

,l>,      -H.-i.  un.vs  o„,,„  .,,,,.,  ^,,,„.  ^,,,,,^  ^1^,,  ^j^^i      ,^^ 

•'■'■■'  ^I'lt  ^".1  s,i,.j  mi::  si,  ,  SI  I.-,  siii;  -  .  -    -> 

-'•!''  -1    1:'  i;I.:J:;  -J^.-jr,  -l.-s  -1.:;!  J,.;;;^  o,;.;.. 


Tahm:  ir 
TiNsioN-  ..i-  .\()iniis   Vai-oi;  i\    Mimphtiks  of  >[ki;iii;v 

,„''";;,.';'''•''";';;   ■''>■   ':^"""";'-;  l"— n.,',   s„l.;K„-t    ni,     run,.    II::.   ,.,„,,.»;.,„„li„j,   ,.,    1; 
''  "M"''.'tiiic  nt   111,.  :,|,    |,„„i  ,1„.  I,:n.in.,.t,i    piessiue  at   Hi,,  tiin,.  „(  1|„.  ,.x  ,.■! 


\p"'t  triHiit : 


Mirildli    Fdll    III  IIUMIMScl    m,s|-|l(\l(iHV    IM  1I\N(, 


I N    M  >  \ 


Tmip.i:   lit 


Mi'n:vrii:K  <'cif;i;i  i  riuNs    to  I{i  i.i  (  i    K'i  mh 


If'    \    Mii:(ii;iM.   Im:(imitii!   vvirir    A 


lli^ 


ScM  i:   III  (I  (' 


SiiMr;irf  till 


i|.l.i|.lillt,-  l|limitiH    ils    fnlMl.l    ill    IllMl'    fl.illl   llio   lll'iyllt    llf    till'    l.tlli.hlrtlT. 


Tin-  llililr   ,»   r.il-  ;i   I.Mi'i tir   «ifll  .1    lirilss  srilli',  illl.l   till'  Miliirs  !,rr  n    litlli'   In 


.■l    tllMIl    f..|    llir  ;;l:i> 


llr 


rill 


;i|'piii\iiii:ilril. 
r.n,|.. 


rill 


IIS    Int     inti'Mli 


■111 


Mill 


'I'mim;  Mil;   Hi  m  i 


1.71 


1.7:1 


L'.!i!l 


■■lilltr   t. 


I>1 


Ml[irl:ttllli 


l.M 


.1-2 


iiliiiiil 


I.S'l 

IMS 
.'Ml 


T.MM.f.  IV 


im;   (i\.-;iiir.s   X'lii.i  \it> 


'I'lir  i.livri\r,|    \nlulih'.   whin    Inillt  I  hlii'ij    \f\    till'   1'ilrl 


Ni'liM  M.    Tl  >'!' 


n:  \ii  i:i'   AMI   |"i;i  SSI  lit: 


I.I    |i|l'isllll'.   will    ^iu'   lllr   M.liHlK 


ll'  tlh 


il  :iir  iriliii'i'il  to  II 


|iMll.liTn;   III   tlir    tl'lil| 


l.l  7r.ll 


r  111*  II 10 


ii; 


l>' 


1:1' 


o,.. 


.Mil 


•  111 

.!iii7 

.:iii| 

.;iii| 

.«*|t7 

.s:ii 

.s;ii 

■'I  J 

vss 

,Ss,~) 

Vs-' 

.s7:i 

IJJ 

.111:1 

.!i|i; 

.:ii:: 

.:iiii 

.:iii7 

.:iiu 

!lli| 

>It7 

.VI 1 

.s:i| 

i:;."i 

.:i.;- 

.IlL's 

.!lj."i 

.'■'-l! 

.!il;i 

.:i|ii 

HI.-, 

.I'lo 

.iMi7 

.:iiii 

117 

.11 1 1 

.!i|| 

.:i.'!s 

.:i.;i 

.'.i.'.l 

.'.ij^ 

;iL'.'i 

■'.'-- 

.Hill 

.;iM 

Hill 

.li.")7 

.11."),". 

.'.i.'iii 

.'.lis 

M\r, 

.!i|ll 

<.i:;ii 

.'.».'.ri 

.!i:'.i) 

.'.•L7 

T.Mll.t 

V 

K.II. 


11,71 


<  Ai.oiiiKs  mil  1  i.iTKi:  Oj 
Ntiiiilicr 


(i.M 


11. s:; 

II.S) 

o.v.-, 

II. si; 

11.^7 

11. <s 

ll.SH 

ii.im 

(>.;i| 

(l.HL' 

11. HI 

n  'Hi 

n.<i7 

n.jis 

n.tiH 

i.nn 

i.»)m; 

l.fi'.lll 

1.71  t 

i.7:;:i 
I.7.-.L' 

l.7<il 

I  7:i-. 

1,;- 

i.Mii 

l.sl:: 

t.sj.-, 

1  ^.;^ 

l.v.-,ll 

i.m;,', 

I.s7.'i 

l.^-«7 

i,:iiiii 
i.:iiL' 

I.HJI 

i.ri:;ii 

l.:l|^ 

I.IHill 

i.:is.-, 

.'  111.". 

.-..ii::  t 

n.niT 

liKI.ATIVK  CAI.liKIKS  CONSIMKIl  AS 

Ciiiliiihv.liaf.'  Flit 
j"'r  I'fiit                       jiiT  (M'lit 

0     ' 


I! 


t 


p 


rr^ 


I  If; 


£   !i  I  H 


i'ii 


1      .    I 


::i1    ■  ( 


?  I 


(  IIAITKi;   I. MM 
SIM;\  ATH'N 

III    lil'li'l-    I"     rillniNli     1|^    \wlli    .■!     >l:||i,|lin|     VMlh     \\\:ir\t     \M'     lli;i\     cnlnli: 

other  c.ill.lili.ilis.    vv,.   sh;ill    lilNl    ..r  .■ill    Mi|,|\     tlic    l.llpiilisiM   ijlllilli;   >l;il', 

tilPll.        A     \;illl.-|ll|c    cll.-lll     <'n||||>ill'll     iViilll     nl)s(   l\:i|ii,||>     IIKKJi'     III     tile    ('■,!<  I. 

irif   Iti>fin:tioii   nl    \Viisliirii:liiii   on    n    umii    ulio    i'm-Ii'i!    Inr   lliirlv-i ,1,, 

is  r.'piddiircil   ill   |"i<;.    |sj. 

The  Excretion  of  Nitrogen.     Whni  .'h.   .•inim.-il   is  si;ii-\c,l.  ji   i,;,.  •. 

Ii\.'  oh  its  own  tisMicv.  1,111  in  (jiiilii;  ^,,  it  s,-u,->  its  pfotiMii,  sn  tlljit  ti 
-•MTctjoii  ,,f  nitfo'j.'ii  t'lilLs  .-iltrr  ;i  tVu  (j.'iv  s  I,,  n  \,,\y  l,.vcl,  lli,.  ,.|i.i- 
(■(■'luiii'iiiiiits  li,.iiiM    ii,,.,iin\|ii|,.  sii|i|.li.'.|,  V,,   hii-  ,is   |.ossilii,..    |-| vt 

'•;lllioiiMll--|I,.     ,111,1      hit,         .\ll||nl|l..-ll     .•|1..;|\>     vlii.-ill     ill     cilllli.irisdll      uitll     I'..- 

till'   stoirs   ,,l'  .•;ii  l„,|n,|i;it,.    v;||\     |si.|rr,-|lil>     i  1 1    . 1  i  iVr  r.'l  i  t    ;i  1 1  i  lil.'l  U.       'I'l:,, 

.■liT  liilii'li  hii-.T  i.  iii.ih  ;iii<|  III,.  li,Tl>i\,,i;i  tkiii  ill  til,'  ,-:iniiv,i|';i.  /' 
iiu,  tl,,    fn-.^l   (,ir  ,/,,.      „/■  s!„rn,li,ni   it    i>  union,   in   tli,'  li.'Hiiv  iiI'm.   t,i  li 

'!'•''      '' MTctioll     ,,|'     ||itliiu,'M     is     ;i,-tllllll\      Lil-c-ltcr     tllNIl     it      u;is     li,'f.. 

^'•■"■^•"   ■"!•   ' 'Ill^''   111''  ioist,.iil   li;|s   I,,', •nil,,.   ,'s|,ililislir,|    in    til,'   tjllM'li- 

o|'  tli.-s,'  ;ii,iiii;ils  .,r  i|s|i,.j  ,-,irlM,|i\,|i;it,.s  jis  tin-  iM.iill  fllrl  liiMt  ,'li;i  I. 
tli.it     v\li,'n    ,-;irlM,li>,|i;ii,s    ;ii-,.    \\  iilili,.|,|,    ,-|s    in    sIm  r\  ;it  inn.    pint, 'ins    :,•■ 

us.',|  III,, IV  ilniii  iiclor,'  ,111, 1  ih,.  i,iir,,._.,.ii  rN,.ivl i.,ii  I mo  yri'iitrr.      W 

\n:\\  s;i\  tli;it  tli,'  li,'i  l,i\  ,,r,,||s  ;iiiiin;i|  l,,.,^  I,,., •,,,,,,.  ,m  i'iii\  .,r,,iis.  'I'll,'  s.i 
tliiiii;-  iii;i,\  o,-,-iir  in  iii;in  \\  ii,.n  th,-  iii,\i,,iiv  ,|i,.|  u;,s  l.'ii',.','l\  ,-,'iil"ii 
illJit,';    tlnis.    ;i!iM,,s|     iii\,ii  i:,|,!\     ill    II,;, II    III,,    iiii|-,„^,.||    ,,i,||,|||     is    l;ii-,.r 

tllf     tllir,|     .111,1      I',, II  I   111     ,i,-|\s    ,,t'    S|;i|    \;,|i,,||      III;,  II      ,,,!     ,j:,,     lii-s,      ;|,|,|      s,.,.,,,|,| 

Ah,itli,'i-    t';i,-t,,r    iiil!ii,-ii,-iii^r    111,,    niip,,.j,.||    , '\, •:•,.(  i,,n    ,|i|iiii.^    tli,.    i;,, 
il;i\s  ,,t'  ill,.  r;is|   is  1 1,1'  :i  III,, II I  it  ,,t'  |)i,'\  i.,11-,  int.-iU,'  ,i|'  nilriMjcii:  tli,'  '.ir,;,'. 

'I'i-  'i'''"  ' II-  'li''  '-Ti'.-it.T  111,.  ..\,-i','ti,iii.      |',y  111,.  s,.\,'i,tli   ,|;iy.  |i,,u,.\,t 

niiii'oi-iii  output   ,,i'  iiiti-,„j,.|i   \\il|   nMi;illy   1„.   i-..;i,.|i,',l   iii-,'sp,'..tivi'  ,.|'  t' 
iii,Ii\i-lii.il's  pi-,,t,.iii  intiik,'.     Diii-ii,;.  III,.  ,^|.,';it,'r  p;irt  of  st;ii\  ,'iti,>ii,  in- 
ol"  tjic  ciM'i'i.'y   n'.piii-,.,!  I,,  i,i;iiiit;iiii   liiv  is  ,|(.|-i\,.,i    iiliu    imI.  ;,s   litil,. 
possilil,'  iMiiiL'  ii,'ii\,',|   tr,,i,i  pi-,,t,'in.     'I'liis  tyii,.  ,,r  iii,'t;il,.,lisiu  L'lsts  iih- 

.■ill    tlu'  ;i\;iiliili|,'    ivs,,iir,-,s   ,,r    |;it    1,;,^.    |„. •\li;iilst...|,    xv  hen    ;i    |||,. 

I'xtciisiv,'  iii('t;ili,,liMii   ,,f  pi',,t,'iii   s,'is  in,   with  tli,'  <',,ns(',pii'ii,-i.  il,;it    ii 

liilrom'i,  rx-r.-tioii  lis.s.  This  is  ir;iiiy  ti,,'  h.-irliili-,-,  u\  ,1,'iilli  ii  is  ,,il. 
cjiHimI  tlic  1,11  iiinrhil  ris,  In  iiilrnij,}!  ,  nnfimr  It  in,|i<-;it,'s  tliiit  all  ll 
onliiijiry  t'lnl  ,.|'  ih,.  ;iiiiiii;,l  ,.,-,,noniy  hiis  l„'i'ii  usci  up.  and  th;if,  it    I 

566 


STAKN  ATlilV 


567 


(iMTiiinai  lUnt'OKv  or  t><{  umicii  las'in/nm  or  aovMCTw.  HtrtN.  nuMnismsi 
METABOLISM  CHART  OF  A  MAN  FASTING  31  DAYS 

APOIL  (4 -MAY  1%.  1912 


•THDIMCMWI 
WIIK  o 


«,VUU<M,r[ldN«.>a. 


MOimWM.fia 


NUT  ni  M  tB-OU 


mTrtMNunM.'c 


leil  l<lll4ISI«IT>aitM>l  ll<I_l4{Jt*>'IIHJM*l 


3  4  »  e  >  »  f  10 1 '  u  i.»  14 1;  i«  n  i,» i»?>ni  tet3i4at«»mta«an 


KiK.     lS2.^^fr..x, 

t  i.n-ti!u  U  •'[ 

i..>!i,    .laia 

"l-l,lll!( 

1     ft 

.Ml      ,i       -l.t 

I  iir    Hav«    of   thr    f 

-T    arc    '^ivi  n 

■ii..,,^:    thr 

i!--i  i---.it 

,    a; 

1     tlu      \. 

ii'-iU^.      ( !■>■  tti    !■. 

(;.    |tt-(ie.|i,t   • 

wii..    t",i«tr.l    f..r    tlilitv   ..M.     .I:i 


i 


i'-i 


if'',' 


i^     > 


Jl^l 


tf-'- 


tf'il 


:.«w 


Ml  'l'\llii|.l>\| 


■I      j 


' ■  ^"''--'O}    t..  Im,,„  tl,..  ^..,•^    ii,s„.,^  tlMiMs.KrH  ii.  ,.,-,|-r  I..  ..I,|..,1. 

^'"^''■""'  '"'■'">   ' ■•''""li"  lilV      \V..,k,.i..'  -iii.ilnl  IriMu'  nil  ^^lmll^l,,! 

iill   MllfMll.t    |>  injl.lr    I..   k,.,.|,   tlii,,,,^   uotU.J    lur  J,    I, til..    In,,.;,.,     lii,.,'    In     Ii,, 

""'^" '■  l->'Nin,..„i   ass..l,.      M,„    ,1,,.,.,  ,,,,.,,,    ^,,   ,,.,„...s..,il..,|   |,^■  ,„•,', 

'""•   ""•    "I    ll"l,.    ,,.„l    x;,h.r    j„    ,vi..|,|„,ir    II,..   ,|,.M,,..|    r,„;us     ImmmiIs.-.    , 
".■     I.JIM,    s,.,.,,,     „|,iv      II     ..alnrirs    „,,.     ,.|\„i|„|,l,,    ,n,;,i„,I     !t.'.!,     ,,l.i„i,„',l,i, 

I  nun   Tills, 

Tll.'v     I'Mrls    ..X|.h,i„     „|,y    ,l„,i„,_,    M.ll-V.lti.,,,     ,1     |„t     „„ v..,...|..s    .I,.,, I 

''""'""•"«'■"    "'•"'    ■■'    !'■• •'II.   Hll.j    «l,:     ||„.    1„|    ,„; „„    ,,,,„|    ,,„,    ,,,,,^^, 

"""  (<><■  ii  l"N..'..r  til,,,.      Tl,..  |.iv,„..,t,-,l  ri>,.  is.  l,.,„.n,.,.,  ,„.,  ,„...^,.„t,.|  I,, 

' '"'"  '"'•  "''''■''  """'''  •"■•■"I  to  ii„l,.-..,t..  tl„if  .|,.„tl,  nun    l„.  ,!,„.  n..t  v, 

•""•■•'  >"  fli>'  .•il.vn..,.  .,r  |-,„.|  ,1,  ,„  „,,,,  „.  „„t,i,„,„.,|  ,|iM,„l,,,i,. f  ,.. 

".•lit,,,!  .,ru,,iis;  ,.  uv,  til,.,-,.   iMM.v   I,..  ,„.  nvnil.-.M,.   „,„t..ii„i   |„  s„,,,,K    ,)„ 

•-'I.UI.U    „f    „|t,.,„,.,|     s....,,.ti,M,     Uill,     tl,..     I,„i|,|i„i;    St ,|„,v     Mlllsl'hav,. 

'  »••.'    I'ilJ.'..    .'iMI  I. 

V.'t    „u\y    is   II,,.,-,.    ,l,i.    ,j„„„,,i    ,.,,i,„_,   „,•   ,„.„,„,,,    ^,,,,.j^^,_,   st„,.vi,ti.,i, 

""  ""'■"  '•"  ■■''^"  •'  'li-'niiiiii.it-  ulil,/„ti f  u|,,.,i   1,;,,  t „s..,|  l,v  ti„ 

•Iil1'''vi,t    „rB..,„s.    .■„•,.,„., I i,,,.,    t.,    ||„.i,.    ,-,.|„tiv,.    „,.tixit.,.s       This    is'v,.,v 

.■l-.'n.l>    sl,„„„  l.y  ..,„„, „„is.,n  ,.r  //„    /„,,  „,   ,..  ;.,!,,  „.,,„.,,  ,,„,,,  ,„.,^„ ; 

■  ',:„„.  ,l„n,u,  sl.,rr„i;,.n.     Th,.  I,  -.rl  „„.l  l„  ,  ,„.  ul,i,.|,  n,„sl   l„.  a.'tiv..  i. 

III.'    IN    t"    Im.    .„.-,il,t;,i„..,l,     |„ ,     , ,     ;;    ,„.,    ,.,.,„     ,„■    ,,,^,j^,    .,,i„j„,| 

^^.•lullt.    u||..,..;,s    tl,..    v„|„„t,.,,N     „„,M.|..s,    tl„.    liv,.,.    ,.,,,,1    ,|„.    ,,,|,,.„    ,„^,. 

•!i.  •••  M,,.!  i;:  |„.,.  ,.,.„!,  ,..s,„,.,i,,.|,     v„  ,|„„,„  ^,„ |.  ,,^;^  ,^^^_,  .^  I 

'"•   '"•.•,.,inl,..|    f.„.  ns   ,1,1,.    t..   tl„.   .lis,,,,,,..,,,.;,,, f    f,.,t.    1,1,1    ,,    s„ni,.i,.„. 

ivn„iii„l..,-  ,-,.,„.,.s..„ts  ,„.„t,.i„  t,,  „,,,U,.  it   ,,|..,i„  that  tl,,.,-,.  „,„st  l,„v,.  I,,,,.,, 
ii  n,„l„li/,.,ii,„i  „r  tliis  s„|,.t;,„....  r,„„,  tiss,„.s  v,l,..,.,.  i,   „;„  „„,  ,,|,s.,|„t..K 

n......ssi,,.y,  sii.l,  ;,s  tl,.'  liv.-r  all.]  v.,hii,tan   „ii,s,.|,.s.  i,,  ....-ans,  sii.-h  as  tl,,- 

li..;,,l.  i„  ul,i.-|,  ,.„..|.,.n   t|.,.ii,sl'„,.n,ali.,i,  is  .„»   ,,„„  „„„  „f  Hf,,      -pj,,.  .j,.,, 
ni-aiis  Ii\,.  al   ll,..  ,.\,.,.„s,.  ,,r  t||,,s,.  „|„,s,.  tiiii,-li„i,s  aiv  ii.MM.ss.,ry 
Til-  energy  output  vw  s,,,,,,,-,.  „,..i..r  .,r  |,..,iy  suifa,  ,■  st..a,lilv  ,'l,.,.|i„.  s 

III    III..    iii,.,i,    ..xanii,,..,!    l.y    h,.,„.,|i,.t.    i|    „as   iCs   ('.    |h.,.   „,, ,.,,,;    „„,,„,.   „, 

■""■'"•"•'■  ••"   " '"'  "'■  "'"■  <!'■•>>   t"-'iil.v-f..iir  li.,ii,.s,  l,„t   „„lv  7:;7  ,,„   ,,„ 

tiN-IVliist  ,la.v  „r  ti„.  sla|.vati„i,  ,„.,i,„i.     Th,"  ..xy-,-,,  intak.'  aii.l  .■a,!.,,,,. 
<li.>\i.|..  ,iui,),,t   ,..iic..s,i.,ii.|i,mlv    ,lii,,iiii,)i 
Til-  l...liavi,„.  „|-  t|„.  nitrcgenous  metabolites  i,,  i!,..  mi,,,,  i^  ,,,•  ,,,,. 

"••"Iiir  iiil..nst,  til,.  f,.ll„„i„..r  ,-a,.,s  Wuvs  ..f  si.n,ili,-a„..,.:    C,-,..,   i.i,,- 

'••■I'i'iv,.h    f;,lls  ai,.|  Ml        X  ,.is,.s.      K,„.  ..,a„,,,|,..  ,„  ,|,.,  |as,  ,lav  ..f  ,V...h„.- 
'!"■   P-iv..|it„y..  „„t|,ut   „)•  Ml        X   i„    ,.,.|ali„i,   t,.  f.tal    „il,„..n.„    „as   :!  1(1 
'Ml   III.,  .iirhll,  .lay  .,!■  til,,  last   it   uas   U.^-.S  i  (•all„.ai.r, .-'     A..i.l.,sis   is  tl,.- 
'•.■Ills...     Til,.  I.,lal  ,.in,.,iint  „r  ..n.atiiiii,,.  aii.l  ..n.atiii,.  sl„.us  „„|v  a  sliulil 

I;, II     1,1(1   ,., Ii,ii|...   ....I... :,...,..   -I..,,  1  ,.         .  *  ~ 

;. '■■•'■'    •'  ■'|'■^<  ..^   .f''..|<-,-,s,-N  aiiu  cn.-itiii.-  ,ii,-,  ,.as,.s  i  ( '.Mt  ii,-;ii-t  ^ 

.'^iiK-o  (•r..atiiH.  is  a  .siil.stii,,,-..  |„.,.,ilia|.  t..  ii,us.-l,.  tiss,,,..  it   is  ,„,ssil,l,-  l,v 


■ 

n 

'! 

i.< 

«TMI\  XTloV 


:.(;!» 


.iM|illliim  tin-  cn'iitilii'  ;iii<l  friMliiiiiic  iiul|iiil  Willi  that  nt  nit  in'.'. mi  t" 
Iniiiiiic  uli.'tlii.r  nil  of  lli.'  iiili.iMrii  lilMrjili.l  li\  llic  liirjik'I'nMi  iil' 
ti>fli'  liiiN  liri"  i\crit(i|.  lie  «|i(lli.'f  Miiiir  ljii>  Imiii  irtailM'"!  cillici-  I'm- 
f    lllll^c•ll•    iIniIi'    iir    li.r     iisi>    cIm'w  liiTc        It'    tin'    iiiiisrlc 


itl 


I  .   -V  111  lll'SIS      111      I  ll 


I  :  .  :iki|ci\\  II    IIS    rMlrlllati'il    I'l 111,.    iTiiilllli'  I'iciil 

III, 111    tliat    raiiiilatril    i'k.iii    tli..    iiiir.....,.|i.    Miillir>is    (.|'    l|i< 

I,  iiiaiiiili'f  tiiiixt  I) iiniiMj;  wiiiT.iis  ir  llii'  liifakilim  II  calriilati  .1  I'nuii 

iiiii.iU.'ii    is   triiMlcr   than    that    i-ali'iilatnl    li 


inn iitiiiil    is   ','ii'ili'f 


IlillliTclltilH' 


iiiii    I'l  I'a;  nil',   c'ti 


itllrT     tl> 


th. 


lan  iiiusi'lf  nnist    \ir  imit  i  ilnilnix  ,      Mnfc.j    ml  i  ipyrii   m-   Ir.'.-   iiiti-i 


L'l  n    in    transit    I'l.nii    |  i>>iic'    to    | 


is^iii-    jiir    iilili/al  inn    k    llif    iiiii-,i 


lik 


HIi'C    of    Slll'h     t\l 


lllll'n'.'ril. 


That  t  lansl'i'i  I'lii'i'  tit'  niti 


rii,'i'iiiiii>  ^l||pslalll■|■s  liiiiii   |ili Ill  plai'c   in  lli 


In 


ti"i:>'n    in   tin'   liiniiil    ainl    ti 


111  stai\atiiiii    is  |ii'ii\i'i|    iji    |,\    the  I'liiistant    ini'srin t'  aiiiinu  ni- 

\'aii    Sl>kr    ;    ami    .  l' i    \<y    tl lI'iTt    uf 

ii's  a  i|i-cii|ri|   iiiriiMsr  ill   I  hi-  ipiit- 


■'|iiiM|s   \witrf  ill  illkill!,'       Till'    lalli'l-  ralh 

It  111'  Mitiii!.'tii.  lull   it  i|iii>  iii't  a|i|iiar  that  tin-  i\tia   nil 


rii'.'ili    1^   iliii'   In 


asiij  |iiiitiMn  liri'aki|ii\\  11       It  is  iimlialili'.  h 


iv\  cv  iM  ,  that 


III  surli   ras 


ll  III'  w  iiiihl  also  III'  an  iiii-n 


lii-rrasi'  III  riiiliiui'iinns  |iriitrin  niflalinlisiii,  siiirc 


ll  '■  \(ashi'i|  iiiit    i'li'i'  nitriiLrtn  uuiihl  ha\i'  In  li 


r-lilai 


Excretion  of  Purines.    Aitl <j\\  at  liisi  t|i,.\   fall 


it.  Ihr  tnia! 


It    iiH'ri'.isi's    as    thi'    I'ast    |prii 


l'irha|is    Ihi'    liisI    iln-li 


11'  til  L'i'nriiil   iisiiiL.'  ii]i  III    li.x  |iiixaiitliiiii'  III' 
tlir  liii'.ikihiu  11  n|'  niii-lfi    I  |iai_'i'  )i.!M. 

Excretion  of  Sulphur.     It   is  iiii|ii.i  ■.■mt   t. 
il|iliiir  ami  iiili-<i'_'.'ii.     In  ll  f  I'lil.v  ila.xs  i.f  >ii 


iiiiisrlf   ami   till'    laid 


i'iiiii|i  111'    till'   I  \i-ii'i|iiii    III' 


il'val  inn  a  ralin  ul    I  i 


X:   1   S 


I  ,  ^  Im'cii    I'liiinii      nit    l.iliT  mil'  ni'    \\ '<    1.   whii-h    is    |ir;h't  ii'.illy    tli,'   saii 


tliat 


Hat    III   iiiiisi'li'    i  1.  !•  .    IJ;    1    .    iiiiliratiii'.'   that    lali'    in    I'astiiii.'   thi'   main 

^"!|l^l'     (if     linitl'in     S|||)|l|\       is     lllllSI'll'. 

.■si'M'i-al  of  tl hanri's  nliscrN  nl   iliiiiiiu  siarv 'itinii    ran    ln'  .-ill  rilmli'l 

'"■  III!'  ninililinii  of  iii-iilti'iis  wliii'h  sii|ii'r\iiii's.  I  I'  ai'iils  air  iji'i'ivi'il 
'"111  iiii'uiii|iii'|i'  I'tiiiihiiNtinii  iif  fat  i -I'l'  p.iu'i'  lis:;',  ami  air  npi  rsi'iili'il 
|.  ;i-iiN,\  lint  >  lii'.  the  aiiinnnl  Irinu  stmirl  inn's  I'DiisiiJi'i  alilf  '|il-l.">  'jTaiiis 
;i  iia\  I,  rs|ii.i-i;ill\  in  iiln-c  imliviiliials  Thi'  larv'c  aiuniniiia  '■•■■,  'timi 
'  ^Miiii'tilin's  2  i.'iaiiis  a  ijayi  is  i'\  iiji-iit  ly  I'nr  Ihi'  |iiii|"isi'  of  la'iil  lali/inu' 
li  .  cNi'css  of  ai'iil.  .Viiolhi-r  (•oiisiM|ncii('('  of  Ihc  .-iriijnsis  is  tin'  ilcrljni' 
'•  till'  aKcolar  li'iisioii  of  »'(».  i  (lairi'  •I-"i4  ' .  :iii'l  it  is  pnssilili'  thai  soini'  of 
!■  I'ii-rnlatiiry  cliaiiiifs  shown  in  tin'  chart  iiia.\  aKo  lie  il  •|ii'nilriit  on 
riic  iiii'lhoii  of  icpciti'il  fastiim  iisiil  t'or  ii'ilnrini;  olu'sitx-  is  (|niti' 
-     ■■  if  till'  aciihsis  is  caii'fnll.v    wati-hril. 

\iaiiy  s)  iiiiiiliirii  vlniinii  s  also  on-iir  in  thr  stai\in'j-  iriianism  Thus, 
Mioiijii/atioii  of  fat  is  ofii'ii  n's|iiinsililt'  lor  a  innnoniifi'il  inrii'asc  in 
lilt  .  onlt'iit  of  Ihi'  hlood    (sir  iiasri'  tiltS  .  aiul  that  of  iinitcin  fxplains 


hhlr 


lli 

i                  ; 

i 

i        -       '                 ' 

li 

!i         .    i 

^iu    ! 


'i 


.-•70 


MITAHiiI.ISM 


tlic  incM'iiir  (.r  Mil  .'iiiiiiiiiit   111'  .'iiiiino  iiitrii!;cii  imt   iiiiU'li   \\v\«\\    ili.it   •.( 
Monii.il  aiiiiiijils    ,\i/..   4   •    :.   [.it    100    c.c.    <,f    l.loodi.     Similarly    v\], 

(■.•iili(ili\.lratfs,   .•ally    in    tli,.   c.iKliii     .   Uw    ],\n,)<l    suu'ar    liociinics    n 1 

'"""'I'  'liiii filial.  Iiiit   llii'ii   iciiiaiiis  stca<ly,     Tliis  is  siirnificaiii    \  !. 

u.'  iviiiciiiImt  tiiat  aft.'i'  two  or  llinv  .lays  .,f  starvation  all  of  tlio  ax.i!- 

"'''•'   i;'.\''"i-'fii   lias  I II   iis,.,l   ii|..      It   in.iical.'s  that    carl.oliydrat..   ,u  ,.- 

'"■  '■^■^'•iiti.'il   I'-.r  lilV,  an.l  tliat   it   is  |,r,M|i |   in  starvation  from   prot.ms 

SIT   |';il;c   (KIT  '. 

Star\;i(inn  ends  in  ,l..;illi  in  an  a.liill  man  in  soiiicw  hat  n\rr  n,,, 
wt'rks  hilt  iiiiii'h  siKJiirr  in  .-liiLlrcii,  liccaiis.'  of  their  niore  active  iii.'i,,l 
olisiii  .\t  the  lime  oCle.-ilh  the  ImmIv  wei-ht  iiia\  he  re.iiiee.j  hy  ."n  ,,.  • 
cent.  The  hnily  1  em  |„.|;|  t  il  Ic  does  not  eliailu'e  until  witlljll  a  I'lW  d.i'- 
of  death,  uhen  il  he-ins  |,,  lall,  and  it  is  undonhte.lly  true  that  if  Ine.ii;, 
are  taken  to  |.re\eiil  eooljii...  of  the  animal  at  this  >ta.,'<',  li^'e  uill  '•. 
pi-ohni'ii'd. 

Death   tVom  starvation   must   he  <li ither  to  a  ".'eiieral   failure  of  ,,!: 

the  eells  or  to  in.jnr.x    of  e,.rtaiii  oiyaiis  that  are  essential   for  lif.v     Sli 

the  loss  of  protein   from   ihe  hody  as  ;.   whole  may   \ai.\    hetween  I'll  .  'M 
.">n  p.n-  cent  .-it  the  time  ,,f  dc.-ith  hy  siaiv  athni.  it  is  unlikely  that  -j.  n./;ii 

failiirr  .•an  |,e  iis  eaiis-.     If  il   were  so.  .|e;iih  w.niM  alwa.vs  iir  w  i..!i 

■- 'i"^'''  ''"-^  "''  piojein  h.nl  oe.'iirrcd.     ( 'erL-iin  orL'ans  evidi  ntlv  .■.■,,„' 

to  i)crlni'm  their  fiiiiclion,  either  h.'canse  they  arc  deprived  of  raw  m,,'. 
rial    lor  llie  dahoration   of  some   sniistanrc    ,  hormone  ■    necessary    for   ! 
or  tieeaiic  the  oi'L-aiis  il  e  n-cUes  war  oiit   from  want   of  noiirishm..nt. 

MiR.M.M.  MKT.Mluj.lsM 

A|.;irl   from  the  pi;ietieal   impoitaih \   knowi.,-  somethiiijj  ah.mt   v  .■ 

'"■'''i^i'"'  "I'  •111  .■'iiini.i!  dnriiiM-  si,-ir\  .ition.  such  kii.ovled.,'e  is  «(  -:•  ' 
\:diie  in  fiiriiishin-  a  st;indard  with  ivhi,-!,  to  comp,-ire  Ihr  ,tul„},„: 
of  ,n<n,Hits  ,niij,r  )i,„„nil  ,  ..,ulil Ioik.  Takiiiu  a-ain  the  iiitro-eli  h.-ikr  ■ 
as  iiidieatinii'  the  eMent  of  protein  near  and  t.'ar  in  the  hody.  I,.,  - 
'•oMsidcr  lirst  of  .Mli  the  .•MiMlitions  nnder  wlihdi  cpiiljhriuni  1u■l^  \: 
-•'''"'"'  "  ""'i''l  !"•  'I'liic  natnral  to  >npp,,se  that,  if  an  amount  ,,i  i  ■• 
tem  eoi.t.'iiiiin-  the  same  auioiiiil  of  nitmireu  as  is  e\ci-ete,l  .hi'  •.' 
starvali.m    were    iriven    to    a    s|;,rvin.j    .iniimil.    the    intake    and    ontpn'      t 

idlro-en  vouM  halaiiee.     We  are  led  lo  make  this  assumption  1 aus. 

know    th,-,f    ;,n^    Inisiness   l,.-,l;il,ee   sheet    shov,  in.j-  .-m    exe.'ss   ,,f  expendr 

"^•■''  '"'■"III i'''l  ''c  met  hy  such  an  ad.iustment.     I'.nt   it  Is  .a  mvv  dr 

cut    inalP'r  with   the  inlni>i<  ,i   h„l,i,i,,    sl,,,t  of  tf,.  l„,d\  ;   f,,r.    if  we 
the  starvinM   animal  .just   eiM.i|..di    protein   to  ,-.,\er  the   nitroMvu    loss, 
shaii  .•ause  til.'  ■■Ncivtion   to  use  I,,  a   total    u  hieh   is  practically  .■,|U.-, 


STAKVATION 


571 


lilt'  starvation  aiimuiit  iilii.t  all  that   we  liavc  yrivcii  as  food;  and  altlioildli 


iV   ( 


lail 


y  jriviiiLr  tliis  anionnt  of  protein  tlicrc  may  lie 


a  sliu;iit  (IccJino  in 


ilic  excretion,  it  "ill  nexci'  eonie  near  to  iieintr  tlie  same  as  tliat  of  the 
intake.     Tlw ly   i^lTeet    of  Midi    I'eedinu;  will    In'   to   iii-olcinir   life    i'oi'  a 


lew    ( 


lay- 


Nitrogenous  Equilibrium. 

,1 


I'll    .itlain    e(|iiililiiium    we    mnst 

nd 


mount  of  itrotein  wliose  nitroLien  eoiitent  is  at  least  two  and  one-Iiall 
limes  tliat  of  tlie  starvation  le\el.  For  a  few  days  followiiii;  tlie  estali- 
iislmient  of  this  |inre  pruteiii  diet,  tln'  nitiiiyi>n  exeretioii  will  lie  far  in 
i-\eess  of  tlie  intake,  hnt  it  will  e;i-,idiiall,\  ili'elirie  until  the  two  praetieall\ 
iiirr('s))ond.  Ilavini:  oni-e  gained  an  ei|nililirium,  we  may  raise  its 
level  hy  <;iadnall.\'   iiieiejisinir  the   protein   intake.      Diirini.'  this  jnou'res- 


■ited  iiroti'in,  it   will  he  fmind,  at   least    in  tl ;.r- 


Nive  i'aisin<r  ol  tlie  inirested  jiiniiin,  it 
iiivora  feat  and  doir),  that  a  eertain  amount  of  iiitroyi-n  is  retained  hy 
llie  body  for  ii  day  or  so  ininiediately  follow  inir  eaidi  inerease  in  Jiro- 
lein  intaki'.     'rh(>  excretion  of  niliniren.  in  other  wcinls.  does  not   imniedi- 

,itely  follow  the  dietetic  increase      'i'lie  ; unt  of  nil  roei'ii  thus  retained  is 

loo  irreat  to  he  accounted  ;is  a  releiitiini  of  disinleeration  products  of 
protein:  it  mnst  therefore  he  clue  to  an  aelual  luiililins:  iiji  of  new  pro- 
tein tissue     that   is.  u'rowtli  of  ninscli-s. 

N'itroirenons  ei|wilil'rium   on   a   jirotein    diet    ah is   ri'adil.\'   attainahle 

Ml  the  eat.  and  less  reaclilx'  ill  the  iloir.  I'-ul  in  man  and  the  herliivoriMis 
Miiimals,  it  is  im]>ossihle  to  y;ive  a  suflicieiiry  ni  pintein  ;ilnne  to  niaintain 
■  iplilihrinm:  there  will  always  he  an  excess  i.f  .■Ncretion  o\  I'r  intake, 
h.deeil  it  scarcely  rcipiires  aii.\    experiment    tn   pmx.-  this,   fiu'   it    is  self- 


ident  when  ^^  e  eonsiijer  that   there  .ire  less 
icooked   lean   meat,   ami    that    tlieie   ari     lew 


tii.-in    WHIH  •'    in  a  )>ound  of 
nid   eat    over   three 


poinids  a  day.  an  amount,  however,  wliich  wmiM  sean-ely  furnish  all  of 
the  re(|Uired  ealorii's.  .\  pelxili  feil  e\eiusi\ely  on  flesh  is  thcfel'ore 
lieinir   partl.\-    starved.    e\en    altliouuh    he    ma>     think    that    he    is    eatiiiL!' 

niifiiitahle  .-iiid  ai-ti\i'.     This  fjiet  has  ;i  prac- 


iliundantly  and  he  <p>> 
th. 


cal  application  in  the  so  ea 
Protein  Sparers.     \'i 


died    lliliiliiiil  I  nil    fur  iihisilil. 


r\-    (lltlereiit     I  esii 


It^ 


■Irales  or  fats  are   ti'cely   i;i\en    with   tlie   ]ir 
XitroL'en   ecpiilihrinm   can   then   he   lee 


d 


are  o':tained  when  e;irholi\-- 
ilein  to  the  starvinir  animal. 
K'li   less  )>ro1ein. 


01     \  er\      till 


that 


>peak   of  fats  .•iiid   eaiiiohydrales  .-is  heiio.;'  •■jifatiht   si>iinr> 


I  arhnhvdrjites  ;ire   much    hetter    protein    sp^niTs   tli.iii    fats;    indeed    they 


'tlicieiit  in  this  I'cirard  tliat  it 


is  now   eonimo 


ilivdrat< 


are   essen 


tial    for   lif. 


iil\  helieved  that  ear- 
that  when  the  food  contains  im 
ace  of  carholixdrates.  a  i>arl  of  the  c.irhoii  of  protein  has  to  he  con- 
ited  inlii  eaih"ii;> 'irate.  'I'lus  iiiipm-taiil  Irtith  is  supj-,,,!  t-'d  1-y  '■,•;- 
■ni-e  derived    from  other  lields  id'  investigation     e.  ■_'.,  the   hehavior  of 


if 


P 


if 

'l 


H 


^If      :i 


III' 


!       I 


i«,        ■■,4 


»: 


-u'2 


mi:taii(ii.ism 


<l''-il'«'ti<-    iMticnts.    ii,    „|,„„i    fli,.    |M.u,.|-   I,,    us,.   .■;,rlM>liv.lnit...s    is    -r,.;,tl 
•I''l'ivss,..|..      Th,.   „„iik,.,|    i>n.t,.ii,-s|.,nii,-  ,-„.|i,m   „|-  .•.•i,l„.l,v,l,i,1.-s  is   II 

'"""■"'■''    " """■'■    "'-'.^       "••'""■l.v,    Uy    tl„.    ,-,,.,    ,1,,,,    „,;,.,,„    .,,.,,.„^ 

'li'MiNisI,    tl,..    iM-ol.i,,    l,n.;,l<,|„«„    ,\unu'^    st.-,,v;,ti„n    hv    frivj,,..    ..,,l„: 

''.■"''■•"'■'•      '"   "''^  "•■'>    "'■  '•""    i'"l I   •v,lih-,.  tl„.  .|,,ilv   Milr ~u   .A,.,, 

'""'    '"   ■■ '    -"'''■''    i'^   •■"M"UMt    in    ,..„n|.!rt,.    sl,,rv;iti,.M.      (•.•„|,„ln 

"ilt,.  st;,,v.-,l,„„    ,s  s.i,|   1„  ,,,t,-,il  .-,    |-,-,il,uv  Ml-  tl,..  „Hls,.|..s  to  ns,.  ;M..-,iM   ',. 

','"■"•  ""■'■• ''^"'    '"■'•'■•''"    '"•  ""•    l"-'"lHHs    u-.iT..   ,.,.,.,,ti,uM    wl,i..irn.s„i, 

"■""•  'li''i''  .l,s,nt..ur;,.i,.n.      At   .mv   r;,t,.  it    hns  I,,.,.,,    fot.n.l  ll,;,t    .•r..,-,t,„.. 
N  ..x.-ivl,.,!   II,  tl„.  ui-i,„.  „|„,„  ,„,  ..;irl„,l,.V(|i-;il(.s  ;in.  ;, viiiiiil)i(.. 

I"  til''  '-.-IS.'  ..r  Mi.-in  lixii.i,  „„  ,.„,  ;,v..f„ir..  ,|i,.t.  ;iltl„.,iirl.  111..  .I.-,iiv  „itrn 

-""  :7'-;"''"-'   i^   ■■ nt    n  ir,-;„„s.   it   ...„    I,..   |,„,„,,,,i   ,„  ,|„„„   ,;   ^,.^„„^ 

';:'"^"'"''    "'•■''    '- '■  ""■   l"-t<'in   tliat    l,;,s  I,...,,    n.,„„v,.,l    f,- ,1„. 

'""•  """""''  '■"'•' >'''•"'■  i^  ^ivn  to  l.finjr  ,i„.  ,,.,.,i  ..,,,„,i,.,  „,,  ,„  ,,,„ 

""•■'.,,■,1  ,i;„iy  n..,..i.-..„„.|,t.      If  ;„,  ,.x,,>s  of  ,•„  il.oli  v.l,-..,t,.  ovn-  tl,,.  ,.„,r.M 
'••■'(""•-■-•"■Nts    IS    uiv,.„.    ,1„.    ,.,.„,..i„     ,„,,    |,„    ,,j|,     ,.,„,,,„,.    ,.^,,1,^^,^.^,    ^^.^, 
'""     '''^""'I'i"^'     'I-     '■MNililH-ium.        It      l,MS     I f„„„.l     tll.U      it      is     „ot     ,1„ 

•'>'""""  •"  ••.•n-lM,l,y.|,-„t,.  Minn,.  tliMt  .l,-f,.r„,in..s  tl„.  ...-.s,.  with   wl.i.-l,  tl... 
'rn.l.r,U,    i.rul.n,    „n,:.„u,n    ,-,u    \n-    n-,,-!,,.,! :   ,|„.   ],Uu]   of  ,„-ot..in    its,.!,' 

'".■.!<-   ,■,    v,.,y    .M-.„f    ,|iir,.,-,. This   h.s    Im.,.„    v,.fv    ,.h.Mflv    shown    l,v 

;"":  '"^ '-"-••"'"■•  "li"  li>M  of  ,11  .|..t,.n „i  |,is  nitfou,.,,  ox,.V,.tion  whil, 

.''',"-    '■^'•'"-'^•■'.^     ""    s'.nvh    .,,,1    .„..,„..    ;,M,|    „ho   th,..,    ,„ .,1    to    s,...    h„u 

'"'."'  ''"'"'■'■'"  "<i'"ls  "I-  l"nt..in  h,.  h.-,.|  to  t.k..  in  o,..h.,.  ,„  i,,in.r  hi,,, 
M't  ,„to  ,„t,„.,,n,:,,s  ,.,|nilil„-i„m.  ||..  fonn,|  th,-,f  h-  h,ul  to  t„k..  tl:.. 
\•'U.,^un•,  .mounts:  ::„  o,„.   „„.„t   ,„.ot,.in.  :!1   u,n.   milk   ,„-ot,.in    :'4  .■„, 

'■"■''  '."■'"'•'"■  -'^  -'"■  I'"''" •"'-•'",  r,4  LMM    I„.;,M   ,.,-ot,.in.  7(1  <:m    h,-,.,-,.| 

'•;■';'""•  ■■""'    '"-   •'"'■    '"'li'-'"- ■"    I"-"l"in.     Th..   o,u^,nis,n    is   .ni.h.ulN 

■■il'ly  t"  s;,.,s,y   ,ts  ,.,„t..i„  .i,.„,an,is  u,,,,-!,   m,.,,.  f,.„|ilv  with   n„.Mt   th;,M 

\\  Itl,    \i.o,.t;,Iil<.    |Udt,.il,s. 

''''^'V'"'''''"'''  '"  ""■  '■ '  '"' '•'•  =  '1''"-"  l'n.t,.i„s.l,.„..„.lsoM  th,.;.- 

""',"••""  ^""'•""•"     >l'.'t    is.   on    ,h,.    |„n,,o,ti„„    nucl    ,.mnn,.r  of   link,.,...- 
"'  "";^"'' -■""!""  .-n-i.isthMt  un,  tol.uiM  up  th..  ,„ol,...uh.      In  uot.", 

'"■'"'■'"^  '"•"  ""•^•:  ' 'li"-^' --  'I'-— ..!|,.,|,  ,,,.,..,,„   in  ..XM...1-. 

""■  -"■"■  l"-l";'""-,  son,..   ,„.o„.i„s  having  .-,    .u-..,,..,,.!,.,.,- o  of  „„..  „' 

••'•""■'"'"•-,.,„.,.„■  ,„„..,,.  J,,.,  ,.  i„,  ,.owof,,„„„„„,..,.,.  ,„;,vl,. 
"•""""'''>  ^"■'•"N-tl.v.i,k...  ,.,.tl,o,„|,  f,„.  ,.,||  of  th..,n  tl„.  sM.n,.  huih' 
niu'  .M,-„..,-,.-,ls  „,.,.,.  .^...i|,|,|„      All,n„„n   ,.,n,|   u'ohuli,,   ,,■,   th..   tnost    in, 

'""'"""  '"■'"■■'"^  '"■  ''' I  ■■""!  'i-n,.s.  s..  ,|,,,t  th..  f ]  must  ..ont.in  ih 

.H-.ss..,ry  ,nnls  f,,,-  tlui,-  .■onst,.u..tio„.      |f  it   f.ils  in  this  ,.....,.,r,I    ,.v,.n  t. 

ti".  ,xt,.„t  of  h„.ki„.  „„i_v  „„,  ,„•  „,„  „„|,^^  „,,.  ,„.,_.^,„i^„,  ^^j„  ;.,,^^^^.  1^ 

iin.ilil..  to  I'lOivt i.ui't    ill...    ...ii'.....    ■■.    !   .1  :',   ii         •  ^. 

p..'..,!..   „i,,i   v.iil    thi'ivi.HV  slin..|.   ffoiii    pjiili; 

M.nvMt.on.  ,„•  ,t   uill   hiiv..  t..  ,.onsti.,„.t    f.„.  its,.|f  this  niissinir  unit.     I' 


Ill 


STAIiVATIuV 


th 


(Mvfnic   illlpillCIlt    llllll    till"    iiMist    \illllill)Ic   pldtcilis    will    1>("   tll(lS( 


tllilt 


(iiitiiiii  iiii  jiiniv  of  units  tlint    i-;i 


H   lie  rciiintiMJ  to   tcifiii  nil   tlic  \;irii'ti 


III'  i)r()tt'iii  ciittMiu!.'  iiitii  till'  slnii-turt'  nl'  the  IkkIv  ])ript('iiis.     Natui'iilly, 
ihc  iH'olciii  wliicii  iiKPst   iicirly  meets  the  re(|uireiiieiits  is  iiie.-it   protein. 


tllilt 


we  !ire  not  surprised  to  (iiid  tluit  less  ot'  this  tliiin  of  iinv  otl 


ler 


protein  has  to  lie  taken  to  irain  nitrou'eii  ((iiiilihriu 


Til 


e  iiK  st   exact    infoi-inatioii   re<;ariliiiir  the  "food   value"  of  ditTeient 


iroteiiis  has  heen  si'ciired  li>  oliser\al  imi ;  on  the  rate  of  i.Md\\tli  of  yoiinu' 
iiiiiiials.      This    method    .\  ields    more    leliaMe    infoi  mr.tioii    than    can    lie 


ured  liv  studies  on  the  nitn 


'llolls  lialaiiee.  I 


leeause  It  is  not  usuallv 


pilSSI 


Me    to    keej)    u|>   the    latter    ohserxati 


nis    foi'    a    siifljeient    iieriod 


lime,  or  to  seeiire  an  adeipiate  iiiimlier  of  data.  I)uriii<r  tri-owth  the 
iiiiildilij;-up  iiroeesses  are  in  exeess  of  the  liieakin'_'-do\\  n,  so  that  the 
rlVeet  is  an  increase  in  hulk  of  the  tissues,  thus  pennitt  iiiu-  lis.  Iiy  the  sim- 
pie  expedient  of  ohserxiiiir  the  liixly  weiirht,  to  ilraw  eoiirliisions  as  to 
the  iiillueiK f  various  foodstiilTs  on  tissue  eonstriletion. 


i) 


i-!l 


1 


m 


I      i   i 


i  r  . 


■f 


*     ! 


i>''    1   "I 


<  IIAI'TKU'   l..\l\ 

M  Ti;i'll<i\   WD  cijdWTII 

III  tli-  uin\Ml,  ,,r  aiiiirKii  tivMi,.>  lu,,  I'.H-tur..  ;iiv  ,.,,ii,M-nH'.|,  ,,ii,.  h,i,,, 
til-  |"'"l"'ily  «\-  tl,..  ,-,.||  ,,,  o,,,„.  II,,  ,i,;.n-lh  f,„tor;  mu\  tli,.  nil„.r.  tl,. 
aviiiliil.ilily  .,r  siiilMl.lr  MKit,.ii;il  t,,  -nnv  upon.  Ihr  fo,.,l  t„rl„r.  Cuinvn, 
insr  th,.  n,.,,ui||  f.-H-lui-  liiil,.  i.  kiM.wn:  il>  viiiial.ility  in  .litr,.|-,'nl  spm- 
nf  .•u,iiii;ii.  it.  iiiv-nlaiit\  , I, .spit,'  |ir..|>rr  a-ljiislinolit  ..f  tlif  f,„„\  fact,,:, 
its  al.ii..i-niali!\  lr;i.lihy  to  tuMi,.i  l,.i  iii.ilioii.  .■t.v,  aiv  .■jll  u.'l|.i<nouii  I,,,' 
.■i(il>arriitl.\    inc'\|ilic-al,!,'    fa, -Is     M,'n.|..|i. 

THE  FOOD  FACTOR  OF  GROWTH 

(iiir  kn,,ul,.,|.^,.  is  ,.,,nsi,inil\    i:i,-i  ,.a>intr  cMM-.Tiiiiiir  tlio  food  fact,.!'  ,  ■ 
Vrnntli.  an, I   rn.Miv    farts  nf  ,\\,vuw  i,ra,-ti.-al   iinportan.-..  liavo  luvn   .v 
|-M.mi!al,„|    in    ,v,.,.,,t    vrars.      In    M..ki,,u.    for   ,1,..    ivlat  io.isliip   of   f„o,|    ' 
vM-outh.    ^u■    must    lirst    of   all    .-onsi,!..,-   uhctli.T    tliis    process   ontaiis 

-'■'■■■"'■'■  '■^"■nWitn.,.  of  cnrr-v  than    is  n. ssafy    tor   ni.Mv  inai.il,.naii. 

II.  ailiilt    lif,..      |,npoi-|,iut    ivsnlts  i„.;irinLr  on   tliis  ,pics|ion  liavo  l.rm 
•■iiiv.l   l,y   nl,s,T\ali.,hs  nn    tl,,'   hasal    iio'tahoiisni    ,,f  >,niii<r  chil.lnn.      I 
-•'■inpiitini.  tlir  .n.iuy  supply  ,,f  fastin-  a-lult  .•iniiii.ils  ..f  .lilTcivnt  si/, 
il    uili   he   ivinnnl.ncl  that    tlie  snialhr  tin,   aniiual.   tin-  jiroater   is   li 
<ii.'i''jy    .■xclianu,'    in    ivlatiousliip    t,,    tlic    ho.jy    u.-i-Iit.    altlnuiirl.    ^^h^ 

'•'"iil'iit-'l    ill   ivl;iti...,s|,ip  t,,  i,n,|y   SUM,-,,-,,   t,,l,..-al.ly   istant    values  ,:• 

ol.taincl       \Vl„n   tli.'  calorie   output    per  s.piaie   nn^tcp   is  .Icteriiiincj 

yrouin-  c!,il,lrc,,,    there  is.  as   » ,■   have  alrea.jy   seen,   clear  evi,l.i 

-reater  en,r,..v    e\p,.n, Inure    ,  see   p.ejv  ,-,-|  I  • ,    p;irti,-ularlv    i„,-,rke,l    in    l„ 

•I"'-'   '"■'■'"■'■  l'"l'"i''.^.     An  increase.}  enerey  nietalH.lisiii  lias  also  I n  •■ 

scrihe.l   in   tl,,,  cas,.  ,,f  infants,  loit   the  un trollal.le  muscular  actixi' 

the  psw-hic  ,|istiirl,an,-es.  etc,  may  e\pl;iiii  the  result.  Kvon  after  ,i 
eouiitiiiL'  th.'se  lactors.  houever,  it  is  p,,ssil.|,.  that  there  may  he  a  r. 
tain  inllueii,,e,  .hpen-lin-  pri.hahly  on  the  a.-liv  mass  of  irroMini:  pn^' 
l-lasmic  tisMie,  uhich  stiinulates  th.'  en.'rey  exi.en.lilure.  The  (piesti 
is  not  >.t   (inally  settl.Ml. 

The  Relationship  of  Proteins  to  Growth  and  Maintenance  of  Life 

Silice    pro',  iii    c.;!i-' it!]'--.    'I-,-    ■■ I-T -•■■•■      i     ■••■■■  •      ■ 

'  ".,.■■    ,i.i     ,, I ,1    ,||,  liii.'ai    oasis   ,,|    llic   cell. 

is   natur.-i!    t.,  .jev,,te   altentimi    in    the   tiist    pla.-e   to   this    f 1   prin.-.p 


SI  iiininN    wii  (iKiiwni 


;»<;> 


I     tlic  pioiiii'i-  iiiv  cstiLMliniis,  stmliis  (111  iIh'  iiiti-Kucn  lialaiice  in  yoiiii'^- 

iiniils  yii'ldcd  icmiMs  li'ipiii  \Uiic!i  it  was  ('iiiii-lnded  that  tlic  PllMditil)ll•^ 

.1    tlic  disiiitciiratioii   ul"  pnitciii   arc   less  developed   in  youiiK  animals 

tiiiii  ill  adults,  so  tliat  tlie  i.'ici\^  iiiir  inuaii-i  rajiidly  withdraw  circiilatiim 

: 'ntciii  aiid  liiiild  it   iiitci  tissue  |)r(itciii. 

Ill  ciiiisidciatiiiii  (if  tlic  a<-eiiiiiiilatiiiii  ul'  data  e\tcMdiin_''  over  several 
>  i-ades,   KiiliMcr  denied   these  eiiiieliisiniis,  anil  sliuwed  that   the  diet    "t 

•  !■  trniw  ini:  infant  is  li>  nu  nnaiis  relatively  ri<'li  in  |)r()tein.  lie  enn- 
.    idecl  that  "irrnwtli  is  not  |)i'iip(ii-ti(inal  tu  the  tpiantity  nf  jn'ntein  in  the 

ii.t."     Inipintaiil   thniieh  this  piiinecr  «ciik  may    have  hecii  in  the  dc- 
rliipincnt  cif  iiMi'  present-day  eipneeption,  tin'  viewpoint  of  the  men  uho 

■    iried   it   oil!    was  very  iiiiieli   narrowed   on   ac iint   of  the  itaiicity  of 

:  iiiiw  ledi;c  coneeiniinr  the  stiiictnre  of  the  protein  iiioIimmiIc.     Xo  allow- 

e  was  made  for  the  fact,  whicli  has  recently  been  lirmly  eslahlished. 

■'i.'it   the  protein   molecule  inn\'   var.v  cNtreinoly   in   rcfiard   to  the  units 

•  •:  which  it  is  c.'iiposcd.  and  that  the  j,'ro\\  in<r  tissues  may  dcniaiul,  not 
^"  Miiieh  an  aliiiinhincc  of  protein  as  such,  hut  rather  a  pr  i]ier  sui)i)ly'  of 
,  n  the  luiildiiiL'  st s  wliich  arc  rciinired    for  'jrowth    i  .Mciidcl  . 


\\  1 1 1  VI  ivi   ( '. i\'i'\i:iNci\  111     \\M\ii    \.  ih-  ( l!;i  ,r,  I  1.  i:i    I  Is  im:ui  s  ,i<  m    l'i;ii,'   i- 
((• i-'iWA  l,y  r.  1'..  ()>l.onio.  llUli* 


— 

"i\.\i.- 

- 

" 

i>.\ 

1   AS    IS- 
II.IMI 

lilMlS 
11.1111 

1.1. 1  \iii\ 
II. nil 

;i:is 

11:  .--!1\ 

Ml. 'Mis 

MISl  1.1 

,.,,,.,. 'I 

0.1111 

:!.sii 

II. ."is 

4.1! 

nine 

1..-11I 

^^.-^- 

l^.i'i' 

l:;..-''.i 

.'l.lill 

^  lis 

s.l 

nil' 

T.lJip 

:;..'iii 

:!.:u 

l.ss 

ii.:;ii 

I 

2.0 

'■nil' 

'.'.:•.,■. 

1(1.71 

<;.<iL' 

l'.l..'l.'l 

1  i.."iii 

S.HM 

It.:; 

1 

li.Ta 

r...")!! 

i.i.i;:; 

'.1.1 1 1 

i.iii 

■I   .to 

S.lt 

nvliil.-iniTii' 

''.l-'e 

"1.07 

ll.  ■•■"'» 

i;.."i."i 

.■;.ii;i 

'I.7.~i 

1..-) 

'.iiniriii'  :iri'] 

l.-|..-,.-| 

!M0 

4:'..r.i: 

I'll.  17 

Is. 71 

T'.sil 

11  Ml 

'.-iitli-   :i.-i.| 

l.:;!> 

:;.:;ii 

|l.."iS 

1.71 

l..'iii 

.'1   ' " 

--.."'l 

iiii' 

ii.."iii 

t 

(i.i:; 

l.ni; 

11..".:; 

ii..'i 

f 

i^ilir 

4..-10 

1.77 

i.r.i 

:'..."i."i 

L'.l.'. 

I..-1.-, 

4.t 

■  .  til' 

f 

4 

c.  (.". 

.' 

l.lili 

t 

* 

'i.lille 

•2.:>i> 

1.71 

l.sl 

11. sj 

■:.]'.' 

L'.IL' 

4..-) 

llliMl' 

.■;.s| 

l.lM 

•J.s  1 

1  ..•..". 

1  1.17 

ll'.lJ 

1 1  ..-1 

'III' 

.■i.ii.'i 

.•1.7ii 

11. '.i:; 

(i.iiii 

i.i;.-i 

!.L':i 

7.li 

i':ii|ili:iiii',  ;il 

ii.Ul        l..'ill 

|iU'S"llt 

l.eii 

1 1.1  II 1 

1  l,>r,ll 

]iii  .-•I'lit 

|iri'sell 

'I'lnill 

I.e.! 

I   '.t 

■"'■-'- 

:;.ii  1 

."'..L'S 

1  .'.•!• 

1.07 

ti.-).l!< 

4S.S.-, 

s.-.lN 

ss  s7 

s  J .  J  S 

.■17.1.; 

liiL'.s7 

'llii-^e    :u!;il>-c>     art-     1  "T;;liii:ah.  i  s     i.l     uli.it     .t;  1  .  .ii     1.     It      1 1  <■     '■! -t     .Ii  It  Tiiiiii.-ilt.-iiv     ul     \:iii 
riu'   t'lijuri-i   fnr   the   iv.i.rc    n  1 1  nt   ;i!i.'iV\  - 1  i^   't    v''.p!  n    .i-i     ii-<iti'I. 


I'lom   the  ;',cconii)aii\  in<;  taiilc   irixiiiLT  the   iicrccntaire   of  the   various 
iiiiio  acids,  etc.,  present  in  eerlnin  pmlcins,  it  w  ill  lie  evident  that  there 
I'  very  marked  v.iri.itioiis  in  the  units  of  which  dilTcrent  proteins  nr 
iii|)osed.     If  any   one  of  these  unils  should  lie  essential  for  growth  an  I 


57fi 


mi;tm»ii,ism 


•' liiiiiiiMti   lie   iiiimIpIc   I.,   mi,' fiic-liiic   tlic   niissinc   unit    |„r   ilMli.   ,t 

is  clciir  tliiit  L'f.iuih  c.iiM  not  |.r.ic.f.|  Ik.wcx  .m-  iniirli  |. lutein  mil  .•(.(ii.n  i. 

'"-  "'""  "• •'""ly  unit  w.'  nii-lit  IVc.l  to  til,,  iiiiiinal.     It   is  iin  ii|,|.li,Mi;  „ 

of  the  i.iw  c.r  th.-  iiiiiiiiiiiiiii,  iiiKl  is  .•iii;;iciu"nis  uilli  tli.-  tiiiliiic  of  u'l...  -ii 
uliu'li  hiis  ]„u<i  liffu  kiKPun  ti.  msiic  \\U,-n  cfrtjiin  iiH.i-jiiiic  siihsijin.  -i 
;iiv  uillilicld  fnini  lli,.  -!•.,„  i„y.  ;ii,i„,;,|.     \  ;|i,.t   ,„i^r|,t   l„,  |„.riV.tly  \..,L 

;in<-iMl  ;is  jll.ltr,.,]  Iiy  .■,MM|.;ili  I  n  of  tlic  llitro;;,,,,  i„t;ik..  iilhl  oiilimi,  ■,,„! 
Vfl    if  it    slionlii    f;iil    to  coiitiiiii   r\cii    one   of   tlie   cssentijil    units   nil. I    1 1  ■• 

"l'-''"'^'"    -^l ''■    !"■   iiie.-ip.-iMc   nf  siipplyiiiLr   tills   unit,   tlini    «oiil,l   ti,. 

liic'    lit'   ili;ii|('(|ii;itc'    fill-  uniw  tli. 

;<■    iiii|>ort,'iiit    fii.'ls    ;ir,.    the    ouieoiiie    of    i jerii    work,    nil, I    i|,,v 

liiiM'  li,','ii  ,'st;ililisli,',|  l,\  olis,-rv;iti,iiis  oii  tli,'  ■:r,i\vtli  of  youii._'  ;iMiiii.i'-, 
iVil  witli  ,'i  ••liMsjiJ  riition"  t,i  \vlii,-li  u,'!','  ;i,l,le,l  iiiixtiires  of  iiiiiiiui  ;i,iN 


■■•':  ''  r""'''  ■'"  '"-  •■"  i"'-'i  I--""'-  I'lM-  111.  s. ,s  |.,,,i,„„  ,,,.i....i, 

';■'-       '■'       '■"""     ■'-     "'■"      ""!•     '' I'.         lA.|.,|,|,a      (,..,„      |.,,l.,y,|l,       I'..      M,  ,;.,,! 


K 


!  i 


or   various   j.nit.ins   « iii,.!,    ,litr,r  e,,iisi,|,'r;il.ly    from    ..lie  ;iu,,lli,.r   in   i^ 
"•"i"''  "I'  111,'   units  eiit.'riii.,'   into   tlieir   iii;ik,'-up.      In   siieli   ,'xp,'rii,i,- 
til,'  perio.js  .Inriii-  uhieli    ;;i-,i«tli    is  oliserveil   must    I..'   |ll',.i,,||^^e,l.  sip. 
a  lr,-ii,si,nf   iii,-r,.,-,s,'  in   u.-iuJit    mielit   .lep,.|i,|  „ier,'ly  on   repair  proees- 

"e.-uirin...-  ii,  tissn.'s  v  lii,-],  liaij  pr,'\  i,insiy  f,,r  s,iiiie  r.';is.,u  I u   lu'oui: 

lielii\\    par. 

Anion-  the  iiioM   'iiiportani  oi.s,'r\ati,iiis  have  l,e,'n  lh,ise  of  Lafayett,' 

.Men, 1,1  an, I  T.  I!    (Kl,,,,,,..-  ,-,11, 1  ,,f  MeCollu,,,  an,!  his  eolh r,'itoi"s.     T 

aiiiiii.ils  .•h,is,.n  for  .M,'n,l,'l  ami  Osl„,riie-s  <'xp,'riiii,'iits  w.'r.'  yoiiii-  wl 
rats.  I.ai'i.','  Iial,'h,'s  ,,f  tli,'s,'  animals  \v,'i','  f'i',1  ,>ii  ;,  l,;isn!  mtion  i-onsistiie- 
pr,ileiii-fr,'i'  milk  M',,iit,'iiiiin,_'-  th,'  iii,ire,'iiii,'  salts,  the  siilmi's,  tr;i,','s 
pr,it,iii.    ami    unkimun    sulistaiii'es    lia\iii-    ;in    iiiip.irt.int    inllueii,','    ■ 


wm  M 


NMTIIITInV    AM>   fJHOWTIt  "iT? 

.'iiiutli  vitainiiics?),  to  wliicli  were  Jiddcd  iiinic  ciirlinliv diiitc,  imrificil 
i:it,  aixl  tlic  i)iiifciii  wIkim'  iiilltitMii-c  on  j:rii\\tli  it  was  dc^ircil  fo  study. 
Tin-  saino  diet  was  fed  at  ri'iiular  iiitiMvals  to  a  nivcii  liatdi  of  rats,  and 
I  hi'  wnV'lit  (it'  I'ai-li  rat  was  periodically  takni,  the  oliservatioii  lieini;  pro- 
Iniij-'ed  until  tlie  aiiiinals  crew  tii  niatiirily  and  jirodneed  yoiiii<r,  and  these 
.i.'.iin  urew  to  inatuiit.v,  reprodneed,  and  sn  on.  Hy  idottin"?  the  re- 
sults in  curves,  with  the  tiirie  jiei-idds  aluny;  the  ahsciisa'  and  tlie  avera<re 
\.  eiillit  of  the  rats  of  ea(di  liatcji  alon<;  the  ordinates,  tlie  extent  of  the 
inlluence  of  a  jriven  diet  on  lln  rum  of  uniirlh  was  nlitained.  A  normal 
'Hive  of  yrowth  is  shown  in  No.  1  of  \"\i;.  is:!.  It  was  olitaincd  from  re- 
sults secui'cd    liy    adtlinir    liheral    anmunts    of   casein    to    the    liasal    diet. 


1-> 

— ' -I- — r        1       1 1 —    1^    1 1- 

111 

:  ® 

/^ 

i*e 

■ 

/ 

M* 

. 

/ 

&U 

• 

*/ 

V>0 

lit 

4 

/ 

*  / 

.(.• 

7 

^ 

-.c 

'  ,^ 

'.  (5) 

/ 

/ 

\ 

J 

AV 

i^ 

?/ 

i^ 

. 

/ 

J' 

^s 

/  <'y 

/^ 

»•»»                           tun  tfiviiion  •  «o  Mfi                Dar<  cacn  diviiion  •sodiyt 

I'iK.    l.''!.      I'urvi-    iif    nr-ntli    c.f    r.il~    .11    l.i-;,l    i.ii:.,!,,    |.|ii*  lln     iir^li  ln<    iii.Iii  .1;.  .1.       In    c:\t\c 

111   Mil-  iir.Tl  ..I    111,-  aiMilinn   ..I    ,,111    I.,   uii   Tiia.li  .111,11.    ,i!l.,«.nu.  ■■!    lli,    |..  ii",,t   i.i"l.iii.    l,i.  l.ill. '1111111. 

■■li'iwii;    .111.1    ill    l\     111.-    iH.-(t    ..I'    111,-    :i. Mill-. 11    ..1    .A-lin,     |i,    .1  .1,11,1,111    ,i,„in    -ill.nv  ^,1,  ,  .       ilnni 
1. My.  Ill-    1:.    M.M.Iil   .111.1   T.    II.   (I,li"rlli-.l 

-iiiiilai-  curves  were  olitaincd  with  hiet.illiiiiiiin  of  milk  iiiid  nvalliiimin 
Hid  ovovitellin  of  eiry:.  reihaps  the  most  interestiii}.'  siilistaiices  cap.ilile 
■r  produciiiir  the  normal  eiiive  of  y:ro\\th  are  certain  of  the  proteins  th;it 
I     15.    (Isliorne    has   succeeded    in    sep.-iratinj;    in    ciyst.'illiiie    form    frum 

.■;.'ctalile  foodstlllTs.  These  are  edestill  ( heiii|iseed  l,  <iloliulill  (sipi.-ish 
-•I'd),  oxcelsiii  (l!r,i/i|  nut),  irlutelin  (iii;ii/e).  uloluiliii  (cuttonsceip. 
■hiteiii  (wheat),  •rly<'iiiin  (soy  licaii),  caniialiin  i  heinitseed'l. 

That  growth  i)i(iceeds  noiiii.illy  with  any  one  of  these  jirolcins  when 
'd  aliiindantly  docs  not.   however,   necessarily   indicate   tli;it   each   eoii- 

liiis  in  afleipiate  i)r(iportion  all  of  the  necessary  units  to  meet  tli(>  i>io- 
■iii  demands  of  crowiii<,'  tissues.  In  the  case  of  casein,  t'or  example. 
'lie  of  the  units,   namely,   •jlycoeoll,   wliieli    is   tlie   simplest    of   .-ill    the 


>rl 


■  m 


i;r 


■lU'  Mr 


liJ;^!, 


578 


MKTAIIOI.ISM 


.-iiniiKi  jii'itls.  i>  ciitiri-lv  iiiissinir,  aiitl  niiiitlii'r,  cysfiiio,  wliich   is  a  mi! 
|>liiir-i'(int.iiriiiiLr  iiniiiii)  ini.l.  is  pri'scrit   only  in  sniiill  imKHiiit.     Tlit'  ,ili 

^»'i <>t"  trl.v«''><'"ll.  liiiui'M'i',  is  not  111'  iiii|i(iitimrc,  lifcaiisi'  tin-  oiiraniMn 

call  liiaiiiil'ai'niri'  it  for  itsrll'  (sec  |«aiji'  (i:!(li.  |ii  the  fase  .if  cvslin,. 
which  the  tissues  can  not  iiiamil'actui-e  thoiiisclves,  the  deficiency  has  •■•< 
he  iiiaih'  lip  for  liy  feeilinn  an  excess  of  casein  so  as  to  cover  tlie  iiei,|-- 
of  the  tissiits  I'lir  tills  amino  acid.  I'.y  so  doini;  a  sii])crai)iindaiicc  >,', 
most  of  tile  otlier  units  will  lie  iiiircsted,  anil  this  superainmdaiice  mil 
entail  the  dcstriution  jiiid  excretion  of  the  useless  amino  acids,  a  prociv,, 
liowever,  which  is  conducted  in  such  a  way  as  to  permit  of  the  utili/atinn, 
liy  the  orijanism.  of  a  part  of  the  eiieiiry  which  the  cast-otT  amino  aii.i> 

'iiiii    i^"'!'  I'a^'c  i1';Ti.      It    is,   ijieivfore.  not  entirely  a   wasteful   pr s. 

When  the  suppl\    ,<{  casein  is  limiteil.  on  the  other  hand,  the  eur\. 

of  frrowth  liecomes  siihnormal.  I aiise  an  insurticienf  supply  of  cyst  in. 

is  therein-  olTered  l  Fi-r.  184).  Similar  results  iiave  heen  olitained  in  lip 
ease  of  edestin,  a  protein  from  licmpseed.  This  contains  an  iiisufliciin.i 
of  the  diamino  acid,  lysine.  Fed  in  aliiindance,  edestin  ilmvc  a  iioriiinl 
curve  of  <:ro\\th,  l.iit  when  fed  in  iiisiil"li<-ient  aniouiit  the  curve  failed  i- 

ascend  properly,  which,   houevcr.   it  coiii.l  he  made  to  do  hy  addiiiir  m 

lysine  to  tile  edestin. 

There  is  a  laru'c  firoiip  of  proteins  which  fail  to  permit  id"  any  {rroutli 
no  matter  in  v  hat  amounts  they  may  lie  added  to  the  liasal  ration.    Tin -.• 
include:  lev'umeliii  i.soy  heani,  viftnin   (vetch),  jrJiadin   (wlieat  or  rye  . 
letrumiii   (pea),   le^'umin    (vetch),  hordeiii   (liarley),  coii<rlutin   (liipiiif 
p'iatine  Mio.  ,i.  zein   (inai/e).  phaseolin   (kidney  liean).     The  adcipiacN 
to  maintain   f;idwtli  of  any  of  these  pure  jirotcins  varies  aecordinu'  '■■ 
the  deficiency  in  their  amino  acids.     In  the  ease  of  cliadiii  of  wheat  "i 
rye,  ulycocoll  is  lacking',  and  lysine  is  present  only  in  small  amount  ( s. . 
tahle).     The  ahseiice  of  ■;lycocoi|  can  not,  however,  as  we  have  alread-. 
seen  in  the  case  of  casein,  explain  the  inadc.piacy  of  filiadin  as  a  foodstmf 
."or -.'row' h  (Curve  II  in  Fi^'.  lS:ii.     It  must  he  the  lysine  that  is  at  fault.   .\ 
still  m..re  <leti.-iciit  iirotcin  is  the  :,iii  of  maize.     With  this  as  the  oip.' 
I)roteiii  a.lded  to  the  liasal  .Met,  the  curve  of  jrrowth  actually  de.seen.l- 
fCiirve  III  of  Fii:.  is:!).  thus  indi.-atin-r  that  the  animal  is  .sfarvinj,'  an.i 
mu.st  .soon   siic.-miili.      The   niissinj;  units   in   this   itrotciu  arc   Klycocoi:. 
lysiiw  aii.l    tr.x  ptopliiine    is..,'   talil..  on    |.a;;e  ."iT.'il,   aii.l    it   is   very   siiriii;' 
cant  that  if  the  latter  two  amino  acids  are  su])plied  alonjj  with  zein.  : 
almost   normal    curve   of   -.'rowth    will    result.      Some   improvement    .•,. 
even  he  lirouj:ht  ahoiit  hy  fiiviiif;  tryptophane  alone;  that  is  to  .say,  fl  • 
curve  assumes  a  horizontal  line  instea<l  of  descen.liiiir,  indieatiiitr  tl^i' 
aUhoufih  iliridfi|!ia1i-  for  ;;rowth,  the  diet   is  n.,u    siilll.-icnt   f..r  the  mail 
tenanee  of  life. 


M'TIMTKIX     \N'I>   OHOWTH 


579 


The  iini)orlaiit  IhcI  lll•lll(lll^l|■iltl•.|  l,\  thcs.'  rxpcrinifiils,  is  that  icr- 
liiin  di<l;t  iirr  iiluiunt,  /„;•  //,,  mnnitf  luinn  of  lift  ullhoiinh  tin  i/  ,iri 
iiiiulniiintf  for  i/roirlh.  In  .•..ntcnnity  with  tills  con.'liisi.m,  it  was  I'ounil 
wlioii  yoiin-.'  wliit.'  rats  wnv  fed  \(it|i  i-liadin  alono  for  piTio-ls  «.f  fiiric  .'x- 


1  IK      I.'- 


l'llnl.,Kr.,|.li« 
but   iilaJfiiuali 
niaintrnancc   ami   grovMli 


al   ,,i 


..,;.;,"_  V".  ,■•■■:■■■»;■■' -  '"   -•'""    '■■"""  ■■■■■•   IIII..1   .In!    .I]. I i,,M   picture^;   cm  a  main- 

'    ZlJ,  "   '"•'^■■'l''-'"-    I'"-   «.."». h.mnl.ll,.    |.„i,„..:    a„.l    .,„   a   .lut    Ihat    «as   inadequate   U.tll 


'ocding    those    in    which    they    should    have    I)oeoiiic    full    grown,    that 

lM>y  ri'iiiaiiicii  ill  an  un>rr(.\vn  stuiitttl  comlition.       The  capacity  to  prow 

■■ad  not,  howt'vor,  bepn  k)st,  for  when  the  gliadin  was  replaced  by  milk. 

I  he  animals  resumed  "fiowth  at  a  very  Rreat  rate.    The  capacity  to  grow 


1 1  .  P 


in 


5h0 


Ml  TAlMlt.ISM 


i:i,i' 


lind  only  liccii  Inhihilnl  l>y  tin-  iriiiiloiuiitc  dicf,  aiiil  llic!'o  was  notliiiif 
rcnll.v  aliiiiiiiiial  alii'ul  tlir  stuiitfil  aiiiriials.  Km-  cxariiiilc,  the  n-prodii-' 
live  riiiicticiii  i|i'Vfli.|H'(l  iKiriiially,  as  was  sliowii  in  flic  case  of  a  ymiii'j 
ffriialc  rat  wlii.-li.  al'tiT  liciiiu'  IV.l  with  Klia<liii  as  tlic  sni.'  inntcin  sii]. 
ply  for  l.'ij  (lavs,  was  iiiat.'il  ami  ihihIucciI  four  yoim-.'.  Altlioiiitli  lli- 
iimtlici-  was  still  iiiaintaiiK'.l  mi  tin-  (iliadiii  diet,  the  youiiir  rats  i>ic 
sciilcd  iioriiial  ".'low  til.  for  tlii-y  wt-rc  liviiij;  „m  tlic  milk  siipplicl  l.y  tU. 

iiiKtlii'i'.  and  this  milk,  1 ansi'  it  cuiitaiiu'd  citlicr  fast'iii  or  smiu-  iitlici 

iioci'ssary  ai-fcssory   factor  (vide  infra),  was  an   adc<|iiatc   food. 

After  removal  from  the  mother,  three  of  these  rats  were  fed  on  an  arti 
li<'ial  diet  of  casein,  edestin  and  the  li.isal  ration,  and  contiiuied  the  mn 
mal  course  of  srfowth.  hut  when  one  of  them  was  |)laeed  on  the  -rliadii 
food  mixture  it  immediately  failed  to  -.-row  inopcrly.  It  would  appeal 
from  these  experiments  that,  of  the  two  amino  acids  that  are  missing'  <>\ 
iIcMcient  in  irliadin  namely,  •rlycocoll  and  lysine  it  must  he  the  lysiii. 
that  is  essential  for  crowlh.  This  very  important  eoneliisicui  was  fiiJK 
eorrohorated  hy  lindin-r  that,  in  ynun<r  rats  stunted  hy  |)revious  ".'liadii' 
feedin-r.  <;rowth  immediately  started  when  lysine  was  added  to  the  di.  t 
and  cease.I  aL'ain  when  the  lysine  was  rcmoveij,  and  so  on,  the  expi'ii 
ments  lieini.'  often  repeated  in  various  inoiliticatiiuis.  .Mendel  ami  (K 
horne  call  attention  to  the  relatively  liiudi  iierccntaire  of  lysine  in  aP 
those  i>roteiiis  that  are  concerned  in  nature  with  the  -.'rowth  of  youh- 
animals:  thus,  it  is  ])resent  in  larire  amounts  in  casein,  lactalliumin  am! 
cfTir  viteliin. 

It  is  i)articularly  in  i)rotein  of  veirctaMc  orii:iii  that  indispensalde  unit, 
are  likely  to  lie  missiiiL'.  the  hest  known  of  these  units  Ipcinjr  the  aroiii.itir 
amino  acids,  tyrosine  and  tryptophane:  the  iliamiiio  aci.l,  lysine;  aii'l 
the  sulphur-containin!.'  acid,  cystine.  Some  animal  i)roteins,  such  ,is 
srelatine,  also  fail  to  contain  aromatic  <;nuips,  ami  are  therefore  utterly 
inade(|uate  .-is  protein    foods. 

That  the  alisence  of  one  or  two  units  shoiihl  render  a  protein  tittcrh 
inca|.,ihle  of  maiiitaininu'  life  su-i-esls  that  a  speeilie  role  may  he  take' 
hy  ceit.iin  amino  a<-ids  in  the  maintenance  of  nutritional  rhythm;  tinis 
they  may  he  necessary  for  the  eiahoration  of  some  hormone  or  other  in 
ternal  secretion  essential  t.i  life,  such  as  epinephrine,  the  a<'live  i.rincipl. 

of  the  suprarenal  irland.     This  is  an  aromatic  sul.sta not   far  reinov.; 

in     its     cli<-iiiicai      stiurtiire      fi tyrosine      (.see      pii;.'e     T.U ) .      It      i 

tlKTcfore  natural  to  suppose  that  the  .il.senee  ..f  the  tryptoi)liane  uiii' 
in  zein  is  tlx-  reason  thai  this  i>rotein  is  incapaMe  of  maintaining;  the  in 
itial  hody  wcifjht. 

In  atla.-kiiiLr  the  i>n,hiem  from  this  acupoint,  Hopkins  and  Will k' 

made  ohservations  on  the  siirrinil  /x  riwl  of  i,,„i)i<i  mivi ;  that  is.  tlh 
peri(><]   durinif   whii-li    the   animals   survived    when    fed    on   a   hasal    diet 


MTKITION    ANI»   (IKIIWTI 


nst 


iiii\r<I  cillici-  with  /i-iii  nloiic  ..r  uilli  /.cin  plus  siiiitll  <|uiiiititii-M  <>(  try|>- 
lii|iliiir(('.     It   was  found   tlml,  with  /ciii  alimi',  tli 


(•    lllll-C    Wl-I'(-    IIIIH 


hie  t( 


tiiaiiitairi  ui'iwth;  thi\v  h.st  in  Wfi«lit  ami  <li(<<l  in  fmin  alunit  a  week  to 
iiliiint  a  innnth.  Olhrr  mice  IVij  nri  the  saiii<>  amount  of  liasal  ili<t  ami 
/(•in.  Iiiit  to  wliiih  WHS  also  aihinl  Home  tiyptopliaiic,  allhoiiifh  tlii-.v  ili<l 
not  tirow,  were  capahjc  of  inaintainirii!  thrii    ImhIv  wciirht   and   lived   in 


sonic  instaniTs  foi-  m-arly  a  month  and  a  half.     There  were  oil 


icT'  nil 


lions  of  the  difl'erenee  in  the  enicienev  of  the  t\ 


di<•t^ 


Tl 


liea- 
fed 


n  the  /ein  alon 


e  were  very   iiiaetive,  and   remained   for 


a  eonsii 


lerali 


period  of  the  time  in  a  eon<liti 


L  es  w  ere  half  eh 


'1. 


if  torpor.     The  hair  was  ruflled,  ih 


d  till 


fe.'t 


d  tail   were 


d.     Tl 


10  niii- 


iiials.  Imwever.  yave  evideiiee  of  havinir  a  Kood  appetite.  <lii  the  otlier 
liiind.  the  miee  to  wliieli  lr.\  ptophane  was  also  iriv<'ii  manifested  a  strik- 
ini.'ly  dilTereiit  hehavior,  heinn  iH'tive  and  more  or  less  normal  until 
.lust  liefore  death.     That   Itoth  m-oiips  of  animals  failed  to  live  more  tl 


■an 


lorly-foiir  or   forty-eiirht  days  is  prohalily  to  lie  ,• iiiited   for  liy   the 

iil'-^'' ■  "I  the  /ein  of  the  other  unit,  lysine.     Had  this  lieeii  added  aloui.' 

with  the  tryptophane  it  is  prolialile,  in  the  li^ht  of  Mendel  and  Osl 


oliservations.  that   the  aniiiiii 


lorne  s 


lild  have  survived  liiiieh  |oiij,'er. 


To  supply  the  missinir  unit,  lieodes  iisiii>r  the  pure  amino  acid,  we 
iiiiifi  niiiilo!)  otiiir  protritis  irliidi  coiittiiii  tin  rniiiinil  ttm'nio  iiclil  (Curve 
III  of  Ki;;.  |S4).  That  mi.vtures  of  protein  foodstiilTs  are  desiraiile  has  loni; 
lieeii  apparent  to  those  who  have  studied  praeti.-al  dieteties.  We  must  eom- 
liiiie  the  iiiisiiitalile  protein  with  others  whieh,  althoiiirh  in  themselves 
perhaps  also  unsuitalile,  yet  furnish  us  with  a  mixture  whieh  eontains  all 
the  essential  units  liotli  for  mainteiianee  and  growth.  As  Mendel  jioints 
out,  these  eonsiderations  simcest  that  w  i"  may  lie  alile  to  utilize  certain 
of  the  low  priced  i>rotein  liy-prodiiets  of  the  cereal,  meat  and  milk  in- 
dustrii's.  The  test  of  the  adcipiacy  of  the  corrected  diet  must,  however, 
he  determined  liy  experiments  of  the  type  which  we  have  Just  descrilied. 
It  is  jirolialily  in  stock-raisiii>:  rather  than  in  connection  with  liiimaii  nu- 
trition that  these  facts  will  prove  of  practical  value:  for,  not  only  is  the  diet 
'<(  man  more  varied,  Itiit  it  eontains  animal  iirotcins  in  wliich  the  deficieii- 
'•ies  are  not  so  common. 

>rost  im|)ortaiit  work  of  this  character  is  iieiii^  eoiidiieted  hy  ArcCol- 
liim  and  his  colljilioralors.'^  It  would  take  lis  heyond  the  coiitiiies  of 
this  hook  to  discuss  the  results  in  detail.  Iiiit  it  may  lie  mentioned  that 
they   have   shown   that,   siiiec   the   ade.piacy   of   the    diet    tlepcnds   on   a 

Miiiltiplieity   of   factors   liesides   the   amiiio-acid   make-up   of   inoteiiis. 

•■■■me  of  wliieh  ue  shall  di.scuss  imnifdiatel) , — very  e.xiciisive  oliserva- 
tions with  various  food  mixtures  must  lie  conducted  over  loiifj  periods 
of  time.  The  nutritive  values  of  the  common  cereals  adde.l  to  a  stand- 
■u-d  diet  that  had  !iroui;ht  the  animals  (rat.s)  to  the  threshold  of  death, 


III 


n 


"1' 


M2 


M(:taiuimhm 


il'.  'i 


I'!,: . 


i 

I     I 


rm  'i  " 


wore  fouiul  1..  Iir  ii>,  follows:    With  foriiiacul  then-  «n«  itiiuie<liate  iefo\ 
ory  nn<l  nipid  Kiowth,  Iiotli  oC  which  himv  also  secuiv.l  in  ••oiiMidi'rabl< 
(h'UMvc  hy  nh.iit  cinl.ryo  nml  I'liiir.-  wheat  krnic! ;  with  lyt'  and  oatM,  ..n 
fhf  otiiir  hand,  tluTt-  was  little  it'  any  iniiiroveiiiciit. 

.Mm-h  work  is,  of  eouise,  yet  to  I.e  done  iMfoi-e  we  ean  dftrnniiie  the 
fxaet  role  whieh  each  nnit  plavs  in  tlie  |>li,\  sioloirie  di'\  elopniciif  oi 
yonnir  aninmls.  I,,  sum  »/;  what  \m  iilreiidy  know,  it  may  he  said  that 
L'l.\eoeoll  is  not  essehtiiii,  sin.'e  it  eiin  I.e  nialinfaet nred  hy  the  anini.i! 
itself:  that  ti-,\  |>to|.hiine  is  essential  for  iiiaiiitenanee,  piohahly  lieeanse 
it    is  rec|niled    for   the    pi'oduelion    of  eertain    essential    holiiiones,    for   the 

niake-n|t  of  wliii'h  in  its  ahseiii'e  othi  r  tissnes  ninsi  I ne  disintoifintcd. 

Icadini:  therefore  to  a  diniinnlion  in  l»«dy  weiuhl;  jind  that  lysiin?  ap 
pears  to   lie  essential    I'or  -niulli       Tissnes   eaii   I.e   maintained   without 

lysine,  hut  they  ean  in.t  -.'i. w.  for  the  sli-.-ht  Ira.-e  uhieh  most   f I  eon- 

tanis  of  this  iinp..i'taiit  amino  a. -id  amy  I.e  snllieient  for  mainteinmee 
pnrposes.  hut  utterly  ina.h'.inate  for  sirowth.  That  the  yomiif  rats  in 
the  experinii'nls  of  Mendel  ..'reu  noiiiially  while  liviin;  on  milk  suppliod 
liy  tlie  stunted  mother  imlieates  that  llie  re,|nisit.-  I.vsine  mnst  have  lieeii 
produced   in   the  nmther's  hody. 

Ill  the  appiie;ition  of  the  foreifoini.'  principles  to  human  dietetics,  it 
is  niidonl.tedjy  safe  to  fo||o.\  Uayliss's  advice  to  lake  care  of  the  cah.. 
lies  and  allow  the  pioteiiis  to  take  e;ire  of  thcmscKes."  lA.r  cxat'iple. 
hi  the  ease  of  milk  the  dedeieney  .,f  evstine  in  its  chief  protein,  casein, 
is  corrected  l.y  the  presence  of  lac-tall.nmin.  ^^hiell.  thouuh  present  in 
only  small  amounts,  contains  sunicient  ipiantilies  of  this  amino  acid  to 
meet  the  deiiniiids  of  Ihi'  s;i'>"  insf  tissue. 

These  ohservations  on  maintenance  and  (fri>\\tli  su<:«esf  very  infrrost 
iiijr  applications  in  connection  with  /A.  uroirtli  of  Imnors.  Is  it  possihj.- 
that  \vc  niiu'lit  letard  the  trrow  Ih  of  tumors  hy  a  diet  that  was  insufticient 
for  irrowth  while  snllieient  for  maintenance.  In  an  expcrimenf  devised 
to  test  this  juoposition  mice  were  fed  on  a  diet  of  starch,  lard.  laet<.se 
and  trlnten  on  which  they  could  merely  maintain  existence  hut  failed 
to  urow.  S(.me  of  these  rats  weri'  inoculated  with  a  rai)i.lly  <;ro\vini.' 
tumor  at  the  sjime  time  as  iinoth.^r  hatch  of  mice  kept  on  normal  diet,  ami 
it  was  found  that  the  tumor  t'rew  much  more  slowly  in  the  stnnfod  mic 
than  in  the  (.thers.     Due  mouse,  for  example,  on  the  restricted  diet  had 

a  scarcely  vjsil.Ie  tumor  .V2  da.\  s  after  the  ii idation.     When  this  mouse. 

however,  was  j.laeed  on  a  m.rnial  .liet  of  l.rea.l.  milk,  etc..  the  lumor 
immediately  l.efian  to  t:row  at  a  very  ureat  rate."  Too  much  importnn.'c 
should  not  he  phieed  on  this  experiment 

Wp  .shall  lu.w-  pass  on  t( nsi.ler  some  of  the  factors  besides  the  pro 

fein  content   which  ha\e  mii   important  l.earinp  on  dietetic  officieney. 


(IIAI'TKK   LXV 


M  TUITION  AM)  <;K0\VTII  itniiiMi 


THE  RELATIONSHIP  OP  OTHER  PACTORS  THAN  PROTEINS 

The  Relationship  of  Carbohydrates.  A><  \\v  have  mi><>ii  cIhiw lietc,  cjii- 
liiiliyilriitt'H  iii'i'  itlinost  I'l-i'tiiiiily  I'ssciitiiil  t'ur  iinriiiiil  iiD'ttiltoliMii.  If  they 
iirt'  not  (rivi'ii  with  tin-  I'immI,  tln-y  iiiiist  Im'  iniiinifiu-tiii'i'il  out  of  prott'in  l>y 
Iht'  orfriiiiisiii  itself.  It  is  not  snrprisinir.  thcrfforc,  lluit  their  iihsenee 
iVoiii  the  (ii(t  of  u'liiwini;  iininnils  slionM  h'ii<l  to  iilmoiinality  in  the 
rnte  of  ^i-owtli.  I 'ei  I  in  I  lists  |iii\e  not  infreiinenlly  insisted  tliiit  one 
liiiiii  of  eai'lioliydrate  is  none  iidviinliiu'eoiis  for  uTouth  tlian  anotiier. 
This  no  (loiilit  in  th<'  main  is  trne.  Imt  the  \\  lioh-  (|nestion  of  aiie(|Uaoy 
l>i'iii>aMy  ilepeiiils  on  tin-  ili(;estil>ility  of  tin'  eai'l>ohyi|i'ate  and  not  upon 
Its  essential  eheniieal  nature.  It  is  likely  that  the  only  earlioliydrate 
reipiired  li\'  the  tissues  is  '.'lueos<'.  Tile  readini'ss  with  whieli  the  enr- 
liohydrate  of  the  food  lieeonies  converted  into  this  MKinosaeidniride  is 
prohahly   the  only  determinant   id'   its  ellieieney   as   food   material. 

The  Relationship  of  Pats.  Altliouu'h  fats  are  an  invariahle  <-onstit- 
Hint  of  |)i'aetieall\'  e\eiy  diet,  it  is  yet  a  dehatahle  ipu'stion  as  to 
uhether  they  are  essential  to  the  maintenanee  of  a  healthy  normal 
i'1'i.'anisn).  hiflienllies  standing;  in  the  way  of  a  solution  of  this  prohlem 
;ii('  tlnit  it  is  not  only  teehnieally  ver\  ditlieult  to  reniox  e  fat  entirely 
rtiim  the  eoinmon  foodstun's,  hut  also  that  the  simple  fats  are  usually 
;issoeiated  with  suhstanees  havini:  similar  soluiiilities  and  physieal 
properties:  namely,  the  lipoids,  phosphatides,  eholesterol,  pijiiuents,  pte. 
Siueo  these  sidistanees  are  present  in  praetienlly  every  eell,  it  is  almost 
'1  itain  that  they  ean  he  mauulaetured  liy  livinir  protojilasin.  Indeed. 
"Nperiniental  e\  idenee  is  not  wantinir  to  show  that  this  is  aetually  the 
'•;ise.  Althouirh  the  eel!  ean  nuiniifaeture  lipoids,  a  .voinitr  animal  ean 
•ippnrently  not  prow  when  these  suhstanees,  as  well  as  simi)le  fat,  are 
'  iitirely  alisent  from  the  diet.  This  has  heeii  shown  hy  feedinii  yoiiiiLr 
iiiiee  on  a  diet  from  whieli  ail  traces  of  I'at  and  lipoids  had  heen  removed 
liy  pxtraelion  with  alcohol  ami  ether  (Stepp)'*.  On  such  a  diet  the  mice 
';'.fd  iinl.\  a  ft^^  v.eiks.  Tlit\N  i-iudd  he  kfli.t  ;i!ue  iiiUcli  ](iiif;er  when 
■-oine  of  the  alcohol-ether  extract  was  mixed  wii  the  diet,  hut  not  so 
^vhen  neutral   fat   instead  of  the  aleohol-ether  extract   was  added.     The 


Il 


:>iH 


MliTAItor.ISM 


iiiMilinii  nf  llic  ,i>li  ,,r  111,.  Ii|>.>ii|  extract  failfd  to  iiiiiintaiii  tlio  iiiicr,  v.. 
Ilial    till'   lacl^iii;;  Milistaiirc  coiiM   not   be   iiii)i-<;anic   in   iiatuic. 

.Mori'  ivo'iit  and  I'sfi'mlcil  olis.Tvations,  lioucvi'i-,  have  sliowii  that  ncntr.ii 
lat    is  also    n.'ci'ssaiy    lor   tlic   a(!t',|uatt'   and    continued    <ri«.\\tli    of   tl., 
animal.     For  a  |ieriod  ot"  two  months  or  so  an  animal  mav,  as  wo  liav 
seen   I'lorii   Osl.orne  and   Mendel's  expel  inn'lits,   >;vii\\    in  apparently   n.n 
iiial    I'.ishion   on  an  artilieial    fat-  and   lipoi.l-free  diet   eoniposed  of  'ea.sei,:, 
eail.oliydrate  and  inoriraiiie  salts,  lint  sooner  or  later  the  j;reat  majiuit 
of  these  animals  heudn   to  show    failure   of  adeipiate   f;ro\vth.     The    in 
ade.|uately   j.'io«inir  ;,nimals  often   manifest   indieations  of  malnutritiup 
other  than  the  failure  to  increase  in  wei-ht ;  for  example,  inllammatioi 
»(  e.ves,   rouirlieninfr  of  the   fur,   etc.      \Vheii   eertain   fat.s  are  added   t- 
the  inadeiiuate  di.'t,  normal  growth  is  iiTimedialely  resume.!.     Fats  pm 
•lucinjf  this  iH)rnial  -rrowth  are  such  as  hutter  fat,  or  the   fat  extraete.i 
from  efrir  yolk,  or  coddiver  oil,  added  to  the  extent  of  ")  i>er  cent  of  tin 
ration.     On  the  other  lian.l,  vcfi.'tahle  oils,  sucli  as  olive  oil  „v  almmM 
oil,   are   inellicient    in    i)romotiri!,'   -rowth.      That    all   oils   or    fats  do   Unt 
snlKice  to  pro<Iuce  s^rowth,  and  that  one  dose  of  an  adcpiate  oil  or  fat  may 
I.e  snllieient  to  stiniidate  it,  indicate  that  somethinj-  other  than  the  men 
prvs.iiee  of  the  comparatively  simi>le   fat   molecule     that    is,  .some  accc 
sory    material     nnist    lie   the  airency   resiionsihle    for  the    <rrowfli. 

This  ci.iiclusion  is  further  suiiported  l.y  the  interestin-r  ol.servation  nf 
.McCollniM    and    Davis   that    ve-etahle   oils  ean    I.e    rendered   cnieient    fm 
-growth    l,y    sh.d^inu'   them    with    a    solution    of   soap   i)reparcd    l.y    com 
plelely  saponifyin-  hutter  fat   with  potas.sinm  hydroxide  in  the  al..sencr 
of  water. 


ACCESSORY  POOD  FACTORS,  VITAMINES 

In    searchimr    for  the   nature   of   the   a s.s<.ry    food    factors,   tlie    im 

portant   ohserv-tions  whieli   have  I.een   made  in   recent  years  concernin'j 

the  so-ealled  vitamin.s  must  1 onsidered.    These  are  sulistances  essenti.il 

in  the  ilict  for  the  proper  maintenance  (.f  nutrition  in  adult  animals. 

The  cxist.'iiee  of  such  sul.stauces  was  su^-ested  l.y  ol.servati.ms  oti 
the  disease  h,  rilxrl.  which  is  caused  l.y  exclusive  feediuf;  on  i.olishe.l 
fice:  that  is,  ..n  rice  from  which  the  |)ericarp  had  l.eeu  removed  l,y  tli. 
proeess  of  polishiu'.'.  When  i)atients  sulTerin«  fmni  this  disease  uer. 
i.'iven  unj.olished  rice,  the  symptoms  immediately  <lisappear.'d.     Furthci 

investigation  of  tl xjo-t   nature  of  these  sul.stauces  was  greatly  facil 

itate.l  l.y  the  diseovery  that  a  similar  condition  is  readily  iridnced  l.\ 
iVedini:  fowls  on  polished  rice.  The  l.ir.ls  deveh.p  a  polyneuritis,  from 
ulin-h,  lion  ever,  they   very  |.romptly  rec.ver  if  some  rice  polishings  or. 


iJkM^ 


M  TKITIDN    AM)   (lUHWTII 


585 


rtfiT  still,  ail  fxtiact  i<{  rice  polisliiii-rs.  is  a.Mi'.l   lo  tiic  polislud  licc 
is  iiiaili'  i(v  nii'aiis  ol'  sli<;litlv  acid  !t!  |)(m-  ct'iil  alci.liol, 


diet.     Tlic  I'xtrai-I 


(I  from  it   l-"iiiii\  ii 


las  siii-cc.'dcil   ill  Ncpaialiii!,'  a   sulistaiifc   in  iTvstal- 


liiir    loiiii   apiiaiviitly    iclatc^d    to   tlic    pyiiiiiidiii.'S,   wlii.-li    it    will    In 
iiiciiilicrcd   arc   a   diaiactciistic   ciiistituciit    iif  the   iiiiclciiis.      I)i.s( 


IS  0.02  to  0.04  u'lii.  of  this  material  i;ivcii  l)V  iiioiitli  wi 


III  cure  the  poiviieiiritis  of  f 


•piatc 


owls  III   Ironi  si.\  to  twchc  hours;  indeed, 


Millie  eases  the  liird  seemed  tpiite  well  after  three  liours. 

>tance  has  also  lieeii   extracted   (\- veast.   milk,   liraii 

and  if   lias  lieeii  called,   for  want    of  a   hett 
It  is  (piite  likely  that  other  dis 


siiiiiiar  sii 
d   1 


I)- 


iiii  anil   liine   niici- 


er  iiaiiic.   vitamim 


seases,  such  as  sciirvv,  iiia\'  also 


lo  the  absence  of  some  \itaiiiii 


he  di 


the  diet 


which  in  the  ease  of  this  particular  disease 
preserved  food,  the  continued  takin<;  of  u  hie| 
Fresh  fruit  and  other  foods  addtd 


some  sulistancc    nam 


woillil  vceiii 


to  he  al 


e  ahseiit   111 


1  is  so  freipieiitly  its 


cause 


oiild  apjH'ar  to  supply  the  necissarv  vitMinine. 


e\eii  in  small  amounts  to  su(di  a  diet 


It  is  not  the  liiixher  animals  alone  tl 


iieh  siihstance 


lat   MilVcr   fioiii  the  want  of  so 


as  vilamine.     ||   has  lieen  showi..  fi 


a    noriiial    artiticial    eultii 
Miiall  amounts,  it   fails  to 


nieiliuiii    IS    inocii 
iow,   whereas  the 


luxuriantly  in  a  iiiediiim  to  whieh  st, 


example,  that,  ^^hen 
with    \e;ist    ill    verv 


same  ipiantity  will   <,'row 


latcd 


<l  1 


leer  wor 


t  has  U 


Vit 


dded. 


imiiie  is  not  of  the  nature  of  a   fer 


iieiit,  since  it   withstands  lieatiii',' 


to   I'-'O     ('.   for  more  than  an  hour.     The  addition   of  yeast   to  dietaries 
that  are  deficient  in  vifamiiies  is  an  excellent  corrective. 

Uetiirninir  now  to  the  accessory  suhstaiiccs  that  seem  to  lie  adhereiif 
to  certain  forms  of  fat.  we  see  at  oiici'  that  they  can  not  lie  exactly 
the  same  as  the  .sii-c;illcd  vitamine  of  l-'unk,  for  they  contain  no  nitroircn. 
There  are,  therefore,  pn.lialily  two  .Mccessory  factors  concerned  in  ade- 
Miiate  irrowfh.  (die  of  these  must  l.e  lu-escnt  in  the  iiniteiii-free  milk 
uhich  serves  as  a  constituent  of  the  hasal  diet  used  in  Oslioriie  and 
.Menders  experiments,  for  we  h.ive  seen  tliat  animals  will  f-row  mi  this 
tor  a  certain  jieriod,  jirovided  the  proper  amino  acids  are  present. 
Later,  liowever,   they   jijiss   into   a   state   wlicie   there    is   no   friowtli   hut 

:idc<piate  maintenance.     If  now    the  other  ; essory   factor  is  added,  jis. 

lor  example,  liiitter  fat  or  a  small  amount   of  milk  itself  (i.e.,  in  place 


■I'  pnitein-free  milki.  then   l'iowiIi   wil 
Mither  of  the  determiii,int> 


le  resumed  at  its  nornial  rati 


iiia\    liecome  curativi 


ir  5,'rowth  when  tlie  liodv  store  of  them  I 


u'comes  ilepii 


Holl 
ted. 


I   are   essential 


.Met 


ollUIII 


uu'Kcsts   that    these   accessory    factors   should    at    i>resent    he   called   tin 
■fat-solulile  A"  and  "  watersolulile  H."     The  latti 


ills 


fat-f 


milk 


aiid  ill  iiianv   oihci 


IS  jireseiit  in  yoast 


Me  same  as 


riiiik 


s  vitamine. 


Th 


al  loiuis,  and  is  pi 


ihl\ 


e  former  is  soluble  in  the  fat  solvent.s, 


r  I 


n  n; 


fiil 


•■- 


1 


586 


METABOUSM 


heiiiff  present  in  most  animal  fats,  l)ut  not  in  all;  for  example,  it  i^ 
absent  from  tlie  fat  surrounding  the  pip's  heart.  By  using  sueh  .1 
nomenelature  it  is  rreojrnized  that  the  sulijeet  is  as  yet  only  in  an  earl\ 
state  of  development. 

We  may  sum  u\^  the  nuiin  faets  of  this  ehapter  by  stating  that  unnill 
Olid  iiiiiiiiti  iiiincf  iirr  iiinrr  than  a  mrrr  prohlrm  of  rnrnjii  xiipphi 
draiittil  tlmt  this  is  siijjicinit.  in  iiiiist  also  have  a  siiitahir  admi.rtiirr  nf 
hiiildiiif/  iniils  of  protein  and  thr  prisnirr  of  r.rtrniifhi  smnH  qitnntitiis- 
of  snnir  iinkiioirii  ornssorit  siilistmirt s.  These  arc  present  in  some  natunil 
foods  liut  not  in  others,  and  some  are  solulile  in  water  and  others  in  fats. 
They  are  found,  for  example,  in  animal  fats  Imt  not  in  those  of  ve<retaM.' 
orifrin.  Motli  fat-  and  wat.r  sojulile  factors  are  i)resent  in  Iar<:e  (|Uanti 
ties  in  milk. 

Hoth  accessory  focnl  factors  are  necessary,  as  is  illustrated  in  tlie  fol 
lowing  summary  of  experiments  rrom  husk's  "Science  of  Nutrition." 
(third  edition V 


Piiiificil    priiti  in 


•!irl».|iv.li:itc 


M';;i'l:ililr    f.'il  '  iiiiii; 
lilltliT    fit 
vcuvlnl.lc   f:it  ■■ 

liiitlcr    (-.it         I        " 


ills  -—  111"  (;i'ii«l1i. 
"  — ;  ml  (jrowtli. 
'       '  vit;iniinos   (.'ircissiiiy 

fai'tiir  ni  =  no  ^roHili. 
"      •  vitiiniini's  nr  yroutli. 


The  Relationship  of  Inorganic  Salts.     I sranic  salts  afe  also  an  es. 

scMlial  iiitrredient  of  the  diet.  McCollum  found  that  young  animals  soon 
i-eased  to  grow  when  fed  on  a  diet  of  corn  and  purified  casein,  hut  that 
rapid  growth  returned  when  a  siiital.le  salt  mixture  was  ad<led.  Oats, 
wheat,  aiul  lieans  have  also  lieen  shown  to  re(|uire  some  a<l,justment  of  their 
ash  content  to  make  them  adcfpiate  for  growth.  Most  of  the  animal  food- 
contain  in  themselves  suflicient  inor<:aiiie  material,  as  is  evidence! 
annuig  other  things  hy  the  ade<|uaey  of  milk  alone  as  diet  for  trrowiiij 
animals  and  the  alihorrence  of  salt  that  is  shown  hy  strictly  earnivorou- 
animals.  In  the  usual  ndxed  diet  of  man  there  is  almost  always  enouu'.: 
inoriranie  material,  the  salt  which  he  aiMs  lieiiig  largely  for  seasoniiii^ 
puri)oses.  When  a  preponderance  of  vegelaMe  food  is  taken,  however, 
the  salt  comes  to  have  a  I'i'al  iliefetic  \alue. 

The  practical  apjilication  of  the  results  of  these  numerous  and  at 
present  somewhat  liewildeiiuL'  oliserviitioiis  to  the  nutrition  of  nian 
and  particularly  to  the  dietetic  treatnu'ut  of  disease,  is  undouhfedl.v 
very  great.  This  is  especially  so  in  infants  and  u'rowiui;  children,  ii^ 
whom  the  correction  of  some  sliirht  inadeipiai'y  in  the  diet  mav  lia\' 
the  most  proufuineed  results,  not  only  on  jrrowth  and  nourishment.  Init 
idso  on  tlie  puuer  of  resistance  ak'aia-ii  disease  aiid  infection.  The  heo" 
fiein!  influence  of  eoddiver  oil,  for  example,  may  depend  on  some  fat 


NUTRITION    AND   OKOWTU 


587 


soluble  accessory  food  factors,  while  the  miraculous  benefit  which 
scorbutic  children  derive  from  the  addition  of  the  juice  of  limes,  lemons, 
itc,  to  the  food  is  undoubtedly  due  to  such  influences.  The  accumu- 
lating mass  of  evidence  as  to  the  faulty  nutrition  in  animals  fed  on 
single  kinds  of  food  that  fail  to  contain  both  kinds  of  fond  factors 
iinphasizos  the  necessity  in  the  dietetic  freatincnt  of  such  diseases  as 
diabetes,  nephritis,  etc.,  of  seein<f  to  it  that  the  diet  is  soinid,  not  only 
ill  calorics,  i)rotein  content,  and  palafability,  but  also  with  I'csrard  to 
the  presence  of  accessory  food   factors. 


t! 


:ii!  r\ 


f    1 


m  >^ 


^^ 


I 


'l 


\  A 


riiAi>Ti:i{  i.wi 

DIKTKTICS 
THE  CALOiilE  REQUIREMENT 

,  '"   ""■  ■■'I'l'li'"' f  tli''  in,|.orlMii1    r.iHs  fl,:,t    liavr  1mt„   n.vh-u...|   v 

""■  ""•""•'■•>""-'  '•''•■'H-is  I..  llM.  >,.),.,„■..  „(  ,li.,,,l,:.  I ,„...,i„„  aris...  „. 

'."  ''""   ''"  "''>   '''•••■'"'i.M.  with  s..i..Mlil„.  ,• un.,.y  just  ..x;„-flv  lunr  ,„,„,. 

.hn„l  sl,u„l,l   h,    l.d,,    „,nhr   r„r„ln<,   c.n.lif,,,,,.   of  Uo.f^h,   orllril.,       I,,     , 

-"■""'•"'  "■■'■^-  "'•  '^"""    "'"'   '!'<■  .•n.MM.nf  „f  r ,1,;,,    „,,  ,,,,|„i,,.  ,„  ,,,  , 

;\'"'"'""""""'    '"    ' ■"""■    ."i.l    .•...>,.,„.(    of    l,.,.|ilv    ..x..,vis..    tllMl     :. 

;"'""  '"•'■'"'•""■'I  ■•"  'li-'  'in..':  -..i.l  .Ik,1,  iC  ,1,..  { 1  supplv  is  i„a ,,„■„, 

"";,"""';    '"■''"'■"    '""-    "'I'    '■••'li   "IT   "nt    V    i MMlitV    i,Ul    in    ,|MMlitv    ,    - 

"-•II.       -llorM's  ,.-,i.s.,  „„,,,    „,„k  Im'sI    wImmi  ti,..v  Mr-,,  well   f,.,l    nii,!   I' ' 

'7'   "•'■■"  ""■>    •"■'■  "«■"   "..rkni.-  is  ; M   .,|,,,.  ,„.I  „„..  tl...  tn.ti,   • 

I ''  '■■■'"  •""   '■'•  "^<'--<iiH.Mt,.,l  in  ,1,,.  , s -.ti,,,,  „f  ..,11  ,|..ostinns  ,. 

"'":'■■"••     ;■'■'"'"•'■ '"-^      -^^    il'-'>'i    l"'i-'i:.-.r   uiii    ,,„l„.r   snir.T    IV,„„    ,| 

';■■""  •■''"'  """■'••^    "'■  -^'''-v^'ti.m  tl,,-n,  .-.ttniii.t  to  ,,..rCnnn  a  i.i,..-..  .,f  u,„ 
fliMt    ;li..    w,.li-,n,.;,nni.,'    Ii..us..u  ilV    L.-n-ailis    s1i,.mI,1    1...    .1.,,,,.    hrf.-.v    s). 
yiv.'s  |,|,„  ;,  ,„.,,1.      Tl„.  s,,i,it   Ml.-.y  1m.  Willi,,;;  lM,t   til..  (I,.sl,  is  u,.„k       IC  1  . 
••"iil'l  I"'  fni>l,.,|.  In.  shnuM  I,.,  f,.,!  liist  ,u,.|  w.„k..,i  .ftn-wanis      li-si,!, 

';"■   ■•""' '    "'■   ""•■'-   '""   ""'••'•    '■^H-tn,s   ,•„•,.    W,.||    k„.,w„    to    i„(l,„.,M...   11, 

'ii'Miiin.l  to,-  fo,;,|     „,„,„.lv.  irnnvtl,  j,,,.!  .•Ii.„.-,t,..     A  vou„^r    .r,„,,  i,,.,  |,., 

"ill  ..tt..,i  .I..,,,;., I.i  „s  „,„.-l,   \{  „o|   ,„„,,.   f 1  ,!,;,„   „.,„1.|  i,|.p..n,-"to  I 

IIS  |„„,,..r  sli;,,v,    tn„i,   .,   ..„„ipM,iso„   of  Ills   ImhIv   w..ii:l,t    will,    tl.Mt    •■ 
li's  s,.,„o,.s:  .-i,,.!.  .,tl,.r  tlu„^-s  l,..i„ur  ,,,„;,|.  i,  i,  „,.||  ,.„„„„  ,,,_.,j  ^^^,  .^, 

""■'""•■'    '"   '•"    """■''    "I...V    lM.„,tily    ..f    ;• I    ,|,„.i„i,    t|„.    .-ol.l    ,lavs   . 

"int..,'   tl,,i,i   ,|,i,-i„^.  tl,..  s„lt|-v   .lays   of  .I„ly   an.!    A,l-„st. 

That    w..  k,i.,w   th..s,.   fa..|s   i„   a   ._'..„..,al   «av.   i„,Ii,.at.s  that   )h..  lii- 
st-|.   to   tak..   ,„    th..   ..xa.-t   .I..t,.r-„ii„ali.„,    of  ,|i..t,.ti..    ,v.|„i,-,.„„.„ts   is   . 

'"."'  '""    '"'"    """■''  '"'•'■^'>    ""'  I>    -xiH'M.ls  „„.|..,.  vaivi,,^.  .-..n.iiti.,. 

"I  ;i.-tiMiy.     Th,s,  as  u,.  l,av..  s..,.„,  ,„..,y  |„.  ,i,„„,  i„.  ,,,,,1,,^,  „„.  j,^,,.^,, 

Ijv.'  t..i-  so,,,..  t„„..  ,„  a  nspifati.,,,  ,.alo..ii,„.t..,-.  s..  that   w,.  „,av  .u.-as,,, 

""■  "■'''"'■ '"'""■     '"'"  ""■  <-"'i--lHsio„s  .|,.aw„    f,..,,,,   n.si.lN  ..f  oLmtn 

ti'iiis   i„a.I..   „,„|,.,-   s„..h   a,ti(i..i.,l   a„.l   u„„s„al   .•.„„lilions  of  livin^r,   ,|: 

-■■'•    '•'••'\"     'I'.Uc   .iiiMI\    lie    i;,is(.|    that    thfv    I,.....!    ,i. 

:il>|.ly   I..  |...,s.„,s   -..i,,^.  al.oiit   tl,..ir  o,..lii,a,.y    ro„ti„(.  „f  Ijf,'..     To  ii,... 

.-,s.s 


blKTlTlfS  589 

iliis  nl).ic('tion  iii'dtlicr  inctlii.<l.  wliich  we  may  <'all  tlio  stallstlcil,  is  avail- 
il>l.'.  Il  coiisisls  ill  t.ikiiiir  the  iivit.is;,.  .lirt  of  a  lar^r,.  iiiiiiiIm'I'  i.t"  iiidi- 
iiliials  and  coiniiariii!,'  tin-  caloric  value  with  tlic  avcra-rc  ainoiiiit  and 
■  pc  lit"  wi.ik  tliat  tlicv  arc  incanwliilc  called  upon  to  luTfoiiii,  and  can 
iHst   lie  used   wlieic  the  diet   is  accurately   known,  as   in   puMie   iiistitii- 

lioiis.  the  army,  llie  navy,  etc     The  total   f I  supplied  is  then  <livid.'d 

l.y  the  nniiilicr  of  in.lividuals.   this  fiiviii!,'  the  jier  cai)ita  coiisiiiiiption. 
(•hviiiusly  some  <ret   more  than   others,  l.iit   when   a  suflicicnt    nuiiilier  of 

individuals   is    iiidniled.   such   errors   Ik me   eliminated    liy   the    law    of 

;i\  erajjcs. 

The  relial.ilit\    of  this  method  is  testified  to  l.y  tlie  remarkalilo  eorre- 

vpoiideiice  in  the  e  dorie  value  of  the  food  < suiiied  l.y  farmers  in  widely 

.lilTcrent  communities; 

K;iri;iiis  in    rmiiircl iciil .'I. Ill) 

"  "     Vi'i-hiiiiif     .".  ('i';,', 

"  "     \.>u    Y.rU    :!'7S.-, 

"  '•     Tl:,lv   :;..-„;.■; 

"  "     Kiiihiii.l    .'UTl 

Avi'i.-ino  ."  "il * 

•l.ii-k:      rii,-    riiiul.iii,,  iit;il    l;.i.  .  .,f    Niitnli'i, 

Till'  :iv('i;i);i'  iiili;il.il:irit  i.f  \;ni.iis  i-itiis: 

f'""'l"ii   2  (;r,.- 

I':iiis    o,„,.. 

M;iiiiili     ■;  III  I 

l\»illi^sl.ci-f o  •.,,  I    * 

••K'lil.Mir.  

Iii.li\i.|ii:il-i  in  .lilTcniit  .■;illiii:;s: 

T':oii,cis'    fMiiiilii.s     (I'.S.A.'l ..  :;..-|i,(1 

Mi'dinniis'   f.iinilic~    i  I'.S.  A.  > :;.(;ii."i 

l*i'iil'csNiMn;i|   inrn 's   f:iiiiiMi^^   il'.S.A.).,  .':  ."I'lO 

Arniv    (f.S.A.)    ■{ ^-,\ 

\;i\.v    (f.S.A.)    .     .    '..'.""    ("'Mist 

!.\i».r.r  ......     r 

In    ^.-N,  i;il.    i'    i«   usiimIK    <'..iii|.nt.,l    iIimI    ;i    niMri 
Uri(;llill^    711   !,;;.   r('i|uiM's   in   i;,|.iii.<: 

L'.r.IKi    f,,r    ;,    sr.l,.Tll:M\     lit',', 

.'l.OdO   I'm-   Ijolit    niiisiiil.'Lf  \VHri<. 
.■l,."iiMi  fnr  niiiliiini    iihi^.-iiImi  wntli. 

I.elMI   iiMil    IlIiWlO.ls    fill     \iV\     IkikI    tnil   I 
;MrKill„|,. 

These  (inures  .-ipiily  to  the  avera-^e  man.  l.iit  in  calculatim.'  the  caloric 

rcipiiremcnts  of  ji  family  or  ; iiimunity  we  mnst   make  allowance  for 

•he  lesser  rei|uiremcnts  of  wom.Mi  and  children.  Sevend  dietitians  have 
"impili'd  t.ahles  showiii-.'  how  m.iny  calories  arc  cxpciiih'd  .•iccordiiur  to 
■■■zv  .Hill  se\.  and  the  (leiiiiaii  aiilhorilics  iiave  recently  tai^  n  these  ll^rures 
■  ind  from  them  c.ilculatetl  .1  ^'cnerali/cd  mean,  which  shows  in  comparison 


ii!.t 


I;  ■ 


I 


i!f  Ik: 


'I.'; 


... ! 


m.'.i  '  I 


r 


!     I 


''^  MKTAnOMSM 

uith  moil  till'  iHivfMliiw  Ihal  .slu.iild  l>o  alUnvc.l  for  women  and  cliil.lr.  i, 
The  litrmvs  a  it  as  follows: 

^!"'  1(111 

\\  (Unrin    ^^n 

Rov   civcr    111.  .  .    .  ,,.',' 

ii'V  14  ii! ....    '..'.". ;,: 

•  '.ill    II  Hi    .       -, 

••hil.l    in  1.! "".'.'. ,';( 

'"'il'l   l''>    .' ,,., 

•••'iM  -■' ::::::::'  ^i; 

n.iM  uri.ir,  L' ;;;  ^...j 

111  .•al.-iilatinsr  tlio  caloii,.  iv.|iiiiviii..|it  of  tli,.  population  as  a  wliol... 
tlir  n.T..ssif.v  of  niakins;  alNmanc,.  for  the  varyiii^r  ,„.,.,ls  of  men.  worn,  n, 
and  ehil.hvn  wo„l.i  ol.vioiisly  make  the  ealeulations  far  too  eom|)lieafr,| 
for  i.raetiea!  j.iiii.om.s.  It  is  neeessary  to  have  a  faetor  l.y  whi<-h  ^^.• 
may  multiply  the  total  population  in  order  to  determine  its  "man  value." 
This  fartor  is  based  on  th,.  relative  proportion  of  men  to  women  an. I 
ehddreii.  and   it  amounts  very  nearly  to  O.:').  i.  p..  three-.piarters  of  tl,.. 

I'.fal  populalion  ^i^s  •'tl ,an  value."    Knowing  tlie  total  population, 

say,  ot  a  rity,  m  ,.  must  therefore  multi|)ly  this  by  Ol't  in  (.rder  t..  ase.r 

tain  for  how  niaii,\    men  doiti;;  i l.rate  museiilar  w.irk  CiOOn  (M    f | 

lias  to  lie  pro\idfd. 

THE  PROTEIN  REQUIREMENT 

The  facts  considered  in  the  previous  two  chapters  lead  to  the  <|uestion 

To  what  extent   may  the  i)i(.portion  of  protein  in  the  diet  he  J'edu i 

with  safety.'  It  is  evident  that  there  must  he  a  minimum  l.elow  whi.L 
every  one  of  th.-  necessary  l.uildiiij;  materials  of  jm.tein  could  mit  I- 
snpplie.l  ill  ade.,uate  amount  to  recoustnict  the  worn-out  tis.sue  protein 

The  extent   to  which  the  protein  content  of  the  diet  of  man  can   !..■ 
lowered  with  safety  .lepeiids  on  s,.v,.ral   factors,  of  which  the  most  iin 
portant  are:  first,  the  nature  of  th,.  pn.teiu;  second,  the  numl.er  of  noi 
protein  calories:  and  third,  the  extent  of  tissue  activity.     Where  so  man 
factors  must  l,e  ta'cn  into  consideration,  the  only  method  l.y  which  tl,. 

actual  miiiiniuni  .an  he  delermined  ( si.sts  in  what  mav  he  called  '-en- 

an.l  try  experiments.'-  Of  the  many  investisations  of  such  a  natm. 
pn.hahly  the  hcst  one  for  us  t<.  consider,  is  that  reeentlv  puhlished  fn.n 
the  Nutrition  I.al.oiatory  .,f  (*openha-en.  The  subject,  an  intelli-'en 
laboratory  servant,  lived  a  perfectly  normal  and  active  life  for  a  perio,! 
nf  five  months  on  a  diet  .,f  potatoes  cooked  with  margarine  and  a  WW- 
onion,  and  eontainintr  4(l()(t  C,  with  a  t..fal  protein  content  of  29  Kran.^ 
DiiniiK  another  period  he  did  outdoor  work  as  a  mason  and  laborer,  a-  ' 
took  5000  C.  daily,  and  :!.".  sjranis  of  protein. 


PIKTKTirs 


r.9l 


It   is  iiiiixtrtaiit  lo  ('(iiitrast  tlicM-  icsiilts  with  tlic  fiillii\vin<r  liascd  mi 
iimnioi]>iil  sfatistii's  ot"  gross  cons\im])tiiiii. 

Mi'Miir-M.  Foiiii  Mtatistii  s 


I'ROrKI.V 

KM' 

r.MtH'lllVlilUTKS 

(■.M,<IRIKS 

;;iii. 

Kill. 

ylll. 

Kiiiiiy'^Itciy 

-^t 

:ii 

Ill 

i;:'.!i4 

Miiiiirh 

IMi 

)i*l 

in:; 

nnu 

I'ariM 

'.IS 

M 

111.-, 

2!I0.! 

LoiiiIdh 

'.IS 

m 

41li 

26fi3 

It  is  pcrljiin  that  man  i-an  Ifad  a  iioniial  cxistoiice  and  I'ciiiaiii  in  }r<><)<l 
ln'alth  on  very  iniifli  less  |inilcin  than  tlic  100  (.'rains  which  statistical 
studies  show  to  hp  tlic  amount  he  actually  takes.  This  discrepancy  be- 
tween the  amount  which  cx|)eriiiieiit  demonstrates  to  lie  adequate  and 
that  which  lialiit  and  custom  <lemaiid.  raises  the  <|uestion  as  to  whether, 
after  all,  our  instincts  may  nnt  have  erred  and  so  made  ns  unnecessaril.\- 
rxtravajrant  in  our  |irotein  intake.  It  has  hcen  sii^'trested  that  such  pro- 
tein extiaxairaiice  will  in  various  \\m>s  have  a  delctcrinus  elTeet  on  the 
iirjranism;  thus,  that  the  cxcittiiry  or'_Mris,  such  as  the  kidneys,  will  lie 
overtaxed  in  eliininatini;  the  unused  amino  acids,  that  the  constant  jires- 
eiiee  of  the.se  liodies  in  excess  in  the  lilood  will  cause  dejjcneration  and 
slui.'f.'isli  iiietalxilisin.  jind  that  the  excess  pidti'in  in  the  intestine  will 
li'ad  to  the  jiroduction  of  ptomaines,  whose  suhsequent  absorption  into 
the  lilood  «ill  cause  toxemic  symiitoins, 

Important  suiijiorl  to  such  \iews  appeared  to  lie  supplied  some  dozen 
years  au'o  l>y  Chittenden,  who  was  able  to  sliow  that  he  himself  and  many 
iilher  jiersons  doiii<r  dilTerent  kinds  of  work  could  be  supported  "U  daily 
.inioiints  of  protein  that  were  not  more  than  from  one-third  to  one-half 
of  the  amount  usually  taken.  Not  only  so.  but  it  was  averred  that  dis- 
tinct improvcmi'iit  was  experienced  in  the  >reneral  sen.se  of  well-bein',' 
;iiid  of  mental  cflicieney  as  a  i-csiilt  of  the  lesser  jirofein  consiiinption. 

Takinir  these  i-esults  as  a  whole,  it  is  (|uile  clear  that  man  can  jjet 
il'iiiir  under  orilinary  conditions  with  mui-li  less  iirotein  than  lie  usually 
'.ikes;  but  that  really  jirov cs  iiotliiii'.r.  for  the  (juestion  is  not  ran  lie  but 
■■liDiild  he,  so  deprive  liimself?  Ai-e  instinct  and  eustoiii  wi-oii'^  and  is 
•  'hittenden  ri(ilif.'  That  is  the  (|iiestioii.  To  answer  it  many  studies  liave 
been  ma<le  of  the  condition  of  iieoplcs  who  for  eeononiie  or  other  rea- 
sons are  eomiielled  to  live  on  less  protein  than  the  average.  Are  these 
people  healthier,  less  prone  to  infections  and  dcfienerative  diseases,  and 
more  efficient  inentally  than  others?  In  such  studies  preat  eare  iinist  be 
exercised  to  see  that  conditions  other  than  diet,  such  as  climate,  exercise. 
'tc,  are  projicrly  allowed  for.  It  would  not  be  I'air,  for  example,  to 
compare  the  mental  and  bodily  condition  of  peoples  livinvr  in  the  tropics 


I 

1 


H 


I 

m 


1 

1'J 


! 


I ' .;  i 


r>f>2 


MIT.MKiI.ISM 


"'"'  ^*''"  '"•< iM|.anilivfly  liitl.-  prnt.-iii.  widi  tlioso  liviii>r  in  t(>nip.-rat 

^;""'''*'    "'' 'Il-llllic    IMIII-Ii    MIMIC.       Aft.'l'    .lic(.UIllillir    all    of    tlicsc    nlli.  : 

f.iituis,  it  li.is  Ihcu  .|Mitc  .•l.-;ii'ly  slic.uii  tli;it.  \vli.'ii  til."  pn.frin  allow.iiir. 
is  n,„l<ri.ilh,  n,h>,;,l.  tlir  ,mm.|,1,.  us  a  u  1,„|,.  arc  l.ss  roluist,  m.-iifally  ih 

'''''■'"•■•  ••""••  i'i>-l<'fi«l  of  iMMim  |,.ss  pi-, t..  tlu-   vory   .liscasfs  xvlii.-h   ml 

ilsii.il|.\    >uppos.-,l   to  1..-  .III..  t„  o^,.|•l,.a.lilll;  of   tlw  oriranisiii   uitli    lis,!... 
.'Mi.'tory  pro.lii.-ls,  an-  ni..r.>  lialil.'  to  sutT.-r  fr.>in  tlicm. 

Tliaf  a  .l.'.'i,!..,!  rcliiction  in  prot.-in  w.ak.iis  fli.-  .l.-f.-ns.-  of  111.-  ..r-,'aii 
isiu  atraiiist  iiif,.,.ti..ii  is  pn.l.al.ly  .in.'  to  tli.'  fa.-f  that  tlic  lliii.ls  ..f  tin 
1-Hly  ii..riiiMlly  .■..iitaiii  a  -irat  vari.'ty  ..f  so-.'all.'.l  aiilil....li..s  that  is, 
of  hiuhly  .•.piiipl.'x  siil.staiK-.'s  that  ar.-  lar^.-ly  pr..tciii  in  iiafiir.'.  Wh.'i! 
ha.'ti'ria.  or  th.'  poiM.ns  i.ro.lii.-.'.j  l,y  tlifin.  .'liter  tli.-  Imly,  th.-y  an-  im.t 

'';'■  '""^  ""•  """ '■  ""•■"•  .l.'f.'us..  snhstan.-.-s  aii.l  .I.'str.,y,.,i  ..r  n.-utrali/.,! 

\..\\   it  is  .'Icar  that  tli.'ir  sh..iil<l  always  U-  a  surplus  of  pr..t.Mii-l.uil.lin- 

niaf.-rials  fn.ii,  «),i..l,  tli..  anlil..Mlifs  may  1 nstruct.-.l.     Such  an  oxccs. 

vill  c.Mistitulc  a  "factor  .,f  safely"  ajjaiiist  .iis.'asc.  Aii.l  there  nre  fa.' 
tors  .,f  saf.'ty  .,f  aiM.tli.'r  nature  to  I,,-  provi.l...|  for,  tw.)  of  whi.-h  we  an 
in  a  p,.sition  to  appr.  -•iat,-.  In  tli.'  first  pla.-...  th.'ie  must  ahvavs  l.e  an 
a.l..<|uate  supply  of  t ry pt„|)haiie.  ..f  lysine,  aii.l  ..f  .•ystiiio,   not"  ..illy  In 

lii.'ef  til.'  har.-  n. ssiti.'s  .,f  the  pr.it. 'in  .•..nstructive  processes  that  s:o  on 

nn.l..r  normal  .■.m.liti.uis.  hut  als..  to  niak.-  f;.,.„l  the  larircr  amount  of 
prot.in  war  an.l  tear  that  -.'iralcr  .l..y;rees  .,f  tissue  aetivitv  will  entail 
Althoiiirh  iiioihrafe  mus.-ular  .■xercis..  ,lo,.s  m.t  app.-ar  t..  cause  auv  im 
ui.'.liat.'  .'..Msiimption  ..f  i.r.,tciii   i  .•arl.ohy.lrat,.  an.l.  later,  fat  heiiiu'  tl.r 
fu.'l  mat.M-ial  us..,l  to  pr...lii<-.>  h\  y.-t  it  .I.m's  thr..w  a  m.-ater  strahi  ..i. 
the  tissu..s  an.l  ..aus..s  a   !.'r..at..r  w.'ar  an.l   t.-ar  ..f  the   ina.-hinerv,  aii-i 

hen.-e  a  .l.-nian.l  f.,r  ni..re  prot.'in-huil.linu'  mat.'iial.     In  the  s.>con.l  pi; 

Ilu'l-e  are  .M.rtaiii  of  th.-  int.'rnal  s.^ivtions  .,f  the  l.o.lv.  s.i.-h  as  ,.piii..p|, 
nil.'  ia.lr..nalimO,  that  ar..  essential  f..r  lif..,  an.l  as  cru.le  mat.-riai. 
lor  th..  maiMifai-tiire  .if  uhi.-h  .-.rtaiii  aiiiin..  aci.is  are  essi-ntial.  Tym 
sin.,  is  ..lie  of  th.'s...  an.l  sin....  pr.,t,.iiis.  as  we  have  se,-n,  .liffer  fr.un'oii. 
another  .piite  ....iisi.leraMy  in  the  am..unt  of  this  aiiiiu..  aci.l  which  th.-. 
e.-ntain,  it  is  a-lvisahle  t,,  i.n.vi.l,.  an  ex.-.'ss,  so  that  an  a.l.'.piate  suppl^ 
of  tyr.tsiii..  may  al\\;iys  he  a\ailal)l.>. 

Tin.  answ.'r  to  on..  „f  th..  most  important  luacti-'al  .|uesfi.ms  in  .li. 
ti'tiis  nam.ly,  "What  pr,.portioii  of  prot..iii  shoiil.l  the  .li.-t  cuitaiii"" 
.lcp..n.ls  ..n  th.'s..  sci..|i1i(i..  princip!,.s.  The  sour.-c  .,f  the  pn.tein  is  tl,. 
important  thinir.  With  animal  prot,.iii  th.-re  is  no  ,l.,ul)t  that  we  ..oiil.! 
•-'.■t  aln„._r  „iti|  ,„,,,,  ,  saf.ty  |,y  takiiii:  .laily  n.-t  more  than  r,()  .,r  Co 
^-rams.  which  is  ahoiit  half  of  what  we  actually  .•..iisiim...  If  the  proleii: 
IS  of  vrnrtahi,  oii;;i„  ai,,!  ,,,.,,t  of  if  .,f  the  (irst  <|Ualit^•,  as  wheat  an.l 
'"•''■■" '"   l"''I'.i'fitions.  iii.„e  sli..ul.l  he  taken  so  as  to  allow  for  th.' 


I'll  TITK  ; 


.".IC! 


<|i'(ici('iwy  ol"  ciMtjiiii  iiiiiino  jic-iils.  Wlicii  \  i'i;<'tiil)|<'  pnitciiis  cit'  llic  scr- 
iiiid  (|ii,ility.  siicli  lis  tlioM'  1)1'  pcis,  Ipi'jiiis.  Ii'iitils,  I'tc.  iiif  iiliiiic  iiviiiliiMi', 
iiiiirli  laijfi'i' aniiHiMts  ;iri'  mi'cssiiiy.  Siirli  in'iitriiis  HIT  iiiiiil('(|iiiitf  in  tin' 
rase  (if  iriiiwiiiir  cliililii'ii  at  Ii'msI,  and  even  in  a<iiills  it  is  iiii<li>iiliti'iliy 
aihisalilf  tliat  oilier  jirdtcins  slimiid  siiiiplfnirnt  tlii'in. 

T((  iiiMirc  safety,  tliiTcfoie.  it  is  aliimst  iiii|(.'ia1  ivc  tliat  tlic  diet  slioiitd 
I'lmtain  jirnliliis  af  riirlinia  aaiin-is.  ](  I'di-  criiniiinii'  reasons  tlic  main 
soiirec  iiiiist  lie  |iioteins  of  ve!.'etalile  origin,  then  Mime  animal  protein,  siieli 
as  is  eontiiiiied  in  milk  or  meat  or  eir'/s,  should  he  added  to  at  least  one  of 
the  daily  meals.  When  |  eas  and  lieaiis  are  mainly  dependei)  on  for  the 
protein  supply,  they  should  le  taken  either  with  milk  or  one  of  its  prep- 
arations, or  with  a  tliiek  vravx-  or  saiiee  made  from  meat  and  eoiitainim.' 

the  (iiicly  mi 1  meat.     This  must   not   In'  strained  ntV.   for  if  it   is.  the 

s.iiiee  w  ill  eontain  only  the  meat  cxtraetives  hnt  not  any  of  the  inolein, 
uliieh  is  eo.iuulati'd  liy  the  Itoiliiiir  water.  .Meat  e\ti;iel,  in  other  words, 
I'liiitains  no  proti'ins;  it  is  not  ;i  food  Imt  nien'ly  ;\  eoiidiment  of  no  irreater 
ilietetie  \;diie  than   te,i  or  eolVee. 

ACCESSORY  FOOD  FACTORS 

Little  iii'ed  lie  aildecl  to  what  has  atreacly  Keen  s,ii(|  rei_'aiiliiiii  this 
sllh.jeet.  'riie  piaetieal  lioint  to  he  remeiidiered  is  th.it  there  ;ire  at  least 
two  aeeessory  factors  coiieeined,  one  of  them  soliihle  in  fat  and  present 
in  aileipiate  aiiioiint  in  luitter  ami  other  animal  fats  Imt  not  in  \CLretalple 
oils,  and  the  otln'r  sohiMe  in  water  aiicl  present  in  wheat.  \ citetaMes, 
fruits,  etc.  ^lilk  (•initains  hoth  of  these  taetors.  so  that  its  ineliKiun  in 
a  diet  is  a  safe-fiiard  not  only  jiirainst  inade(|iiac-y  in  siiitaMe  protein.  Imt 
also  aijiiinst  the  jiliseiiee  of  aeeessory  food  t'aetois.  'I'lieic  is  little  ilaii!.'!'!' 
of  the  diet  lieinu;  inadeipiate  with  reu:ard  to  fnod  f.ietors  if  ii  contains 
some  fruits  or  u'reen  venetahles  or  nnheated  iVesh  milk.  The  fooil  I'ae- 
tors  are  destidye(l  hy  jirolonired  cookinir. 


DIGESTIBILITY  AND  PALATABILITY 

We  liavc  seen  that  pi-acti<';il  dii'telics  depends  on  se\er;d  factors,  the 
exai't  relative  importance  of  whii-h  can  not  perhaps  he  •.'aULred  in  every 
ease.  Imt  piejiaration  id'  the  lood  so  as  to  m.ike  it  aiipeti/ini;  must  iiii- 
douliledly  rank  hiifli.  The  importance  of  iiniiil  riKil.iini  will  now  he  aji- 
pai-ent.  It  is  the  act  of  makin<,'  food  appeti/inj;  and  thei'cfore  diirestilile. 
It  is  really  the  first  sta^'e  in  dij.'estion,  the  sla<;e  that  we  can  control,  and 
one  tliercforc  to  which  nmch  jil  lent  ion  iniist  he  jrivrn.  espcciallv  when  ii 
hecomes  necessary  to  in. ike  attractive  articles  of  diet  ordinarily  considered 
common  and  cheap.     .Most   people  can  cook  a  laiiili  chop  so  as  to  make  it 


Ml 


i! 


IM 


4SN: 


;(  :; 


H      ;  ! 


.V>4 


Ml  'r\ii(ti.isM 


r.Ms.„MMN  ;inM.t,/,Mu'.  hut  t..«  .•„,.  t,.kr  ih..  rl,..,,,,,.,.  ...it^  of  m.-i.t  ninl  

^'•'•<  'l"M,  H,t.,  ,..M.k,..|  .|,sl...,  i|,„i  „r-..  i.s  |.o,„.li„-  nii.l  i,ttnictiv...     A.mI  tl,.  ,. 

""■  ^""  '""'■'■  "' ■"'  '•'•<'•  ""•  l-'l  "V'--  nn.l  Iri.nmiii^'s  i.n.l  ....r.v-rt  i|„  ,„ 

'-'  tin.  s.,n..  «,n,     Tl,i>  is  tli-  ,vm|  ,,t  .,i  rnokin^r.  m,.l  too  Mni.-I,  n, ,„■.. 

"""'  •• '  '"■  ■■""■"  '"  " "■"•■'  "l.i.l r  r„.,kH,i!  ..xiMTts  mv  n..kn,- 

'",^'"'"   ' I'''-  '•""   ""•-•  "'i'll-'-  '•^'"  I ' Tli-  uast,.  of  a f I  „. 

.•I  l.'ii;.'!'  cilv   is  tnily  ;i|>|)iilliii',' 

•■-".kill..'    l,J,s    oIImt    lMl\ilMl;iL..    -    th.'IM     riNlkillir    Uu-     foo.l    lippcti/ill-         Tl:. 

l"Nit    l,M,s,.„s    il„.    ,„i,s..|,.    1,1.,,,.,   ,„•   ,i„.    Miriil    so    lliiit    it    is    inor,.  ",.,.„,, I 

llMsII,-;,t,.|;     ,t    .|..stn.Vs    , ,  „OI...M„isM.S    illl.l    |.Hn,sit.>S    ill    thc    l.l.'at  ;    it    ,|. 

strovs  .,„tilM.,|i,.s  uhi.-h  M,iL..|,i   i„t,.rtVr.-  uitli  tl..-  ...•tjo,,  of  tl...  .li-rstiv, 
•■•iiH.nis      Tims,  n„i,...„,..,|  ,.,,,   „,„,..  „,.,„_,^,  j,^  „„,  ,|ijr,.st..,|  in  tl...  st.„„ 

■■'"''   ' ""-''  "   '••''""  ""•■  "'■  'Ik'  aMtil.o,|i..s  ulii.-h  p.-..v..|it   tl...  |„.|,sM 

"■'""  "'•""-  '■■  '""   ''"i'"'!  <•"-'  "l.it-.  if  |.r riv  ..|h.«...1.  .-a |, 

•-'.-i.-l.  ;,M.|  V,  in, ,,„„.,  Ih..  ,...-  uliit..  into  a  foam  partly  .i.-strovs  tli.  ,„ 
liiliitiiiL;  siilisiaiici', 

'■■■'""•  ' ''"''"^-  -■""•'hint!  s ihl   h..  sai.l  ah.Hit    .h.-  la.ralln    ,,„„l, 

/"v  u,   i„.,.l.  r,„.  i,   is  „|-,,.„  i„  ,|,i,  piirtirnlar  ah-nr  that  on.,  f is  mm.,.. 

sat,sta,-..,ry  ,|,an  an.-ll.T.  A  ,ii,.t  „f  ,„,,„.  ,„i|k,  ,.._,j,,  ,,„.|  „,,;,,,  ,„.,,.^^|  ^^ 
ai't  I.,  h..  nni.h.vsinl,,.^,..  Im.,.,.,,|s..  t|„.,.,.  js  nothing'  in  it  to  a.-t  as  what  1,;,^ 
I"-"    'mH-.I    ini.stinal    hallasi;   i|,,.,|    is.    a    .nat...ial    uhh-h    will    k.-,.,,  tl  ■ 

""'■""""^  sMtlh-i.nilv  tl I  tn  siian.lat..  th..i,.  nius.Milar  ,n..v..,n..nts      Tin. 

I'.-ill.i-t    IS   JHs!    t„,nis|,...|   in   th..  shap..  of  ....Ih.h.s...   th.-   most    in.poH;,,,' 

'■'"""""•'"  '"■  -"•'■"  '■ '      '''■••'•>.  l'"-'«"s.  .-ahhas:...  saln.l,  an.l  tnanv  fniiN 

.■sp,...,..,i|>   appl.s.  shonl.l  always  , upy  a  pla.-,.  in  th.'  .lailv  im-iiu       \h 

"""■'■  ^•■'l".'''l"  '■ I  >iH.lin.,^  this  l.allast  is  th..  ont..,-  irrain  ..V  win-at    oats 

H-  So  MH,-h  ninst  „ut  l...  tak..n  as  to  pn„|„..,.  a  .-..nstant  intostinni 
"■'•i'i"i"M.  ihhI  ..a.-h  prison   innsl   .{..trrniin..  for  hinis..lf  w  h.-lr  this  linir 

'"■"      ''''"•  'li'1'»'''' .inioni;  various  l.r.'a.ls  is  almost  .Mifir.-Iv  in  tho  -h 

'-'I '"  "hi.-h  lli..y  suppiv    liallasf. 

'I'll.'  all-importani   sul-L.-t  of  foo.l  ,...,.„.mii..s  .-an   r..o(.ivo  n..  att..nti.. 
li'i-'.  -N.-.pt  'n  point  out  thai  it  is  .,n..  whi.-h  must  ho  m..st  .•aivfully  .•„. 

^"'""■■'   '"   "'"■  ■^'■1'"' '■  -ill  l»-nl,l..ms  of  ,li..t,.ti,-s.     An  a.lmirahl,.  a. 

-•oMiit  nf  t|„.  siih.i..,-!  u,ll  l„.  fonn.l  in  (iraham  Lusk 's  ".S.-irii.-,.  of  Nut, 
tion"     thir.l  ...liiion.   an>i   in   .M..Killo|.'s  "Foo.l   Valu«'s."'« 


rilAI'TKI.'  LWII 


Till-:   .MKTAI'.nMs.M   u|'   l'|?'"Ti:iV 


Introductory.     Tl 

'I'll  'Ik'  I' I   "iis  liiiiilL'lit    iiitii  ;i   siilulil liilitiiiii   li\    till'  (liu't'st 


II'   lil'lrr-    |)||\  sh     .iuist> 


I  hilt   tlir   lil'dti'ili   tilki'll 


ive  c'li- 

/viii.'s,  iiliil  lli.it   it   -MiN  thru  iilisdilxMl   into  tl..'  Ill I  iiikI  ilii'fi-tly  ilinir- 

IMir.'itcil  with  tilt'  lixsiirs,  Thf  .liscivcry  <<(  tlic  rii/Miics  Inpsiii  jiikI 
I'l't'lisiii  aii<l  111'  I'l'i'c  .'iiiiiiin  .icicU  ill  ilir  i.','i>tniiiili'Nliiiiil  I'lmti'iils  clcili-lv 
sliou.'il  thill  this  siiii|ilc  •lii'i.i-y  1.1'  Lii'l.iir  cniilil   nut   I orri'ct.     It   w;is, 

fllltllt'lllllPl'i',    I'lilllllj   that    wlll'll   JIM   I'Nci'NS  111'   |i|ii|t'ilis  sllch    lis   I'uu:  iilllUlllill 

'.'iiiiis  ciitrv  to  till'  hlo.MJ,  pjiit  III'  III.'  pii.t.'iii  iippf.'irs  in  an  iiiicliaiiU''il 
.■.iiKlifitin  ill  th.'  mi  1 1.':  ami  tli.il  enzv  mi's  i-apaM.-  ..f  .li..'.'s|  in._r  this  pint. 'in 
I'lit  iii.t  (iIIht  \aiiiiifs  mak.'  tlii'ir  appcaraii.-.    in  tlic  lih....!. 

Al'li'i-  thf   iiijcciiiiii   lit"   f(iici!.'ii   pint,  ins   into   the   Miiinl,   s\  niptoiiis  i>i' 

•  iniiifr   si'vciity    ..tlcii    iJ.'M'l.ip.    rn.ni    th.-    al st    iiistaiitiilicnis    <!cath 

proilnci'd  hy  snake  m'Iimmi  t.i  th.-  slow  I;,  iii'\  ilopin<_^  aiiapli.v  lactic  I'cai- 
lions  «hicli  I'lillim  the  inj.'etion  into  tli.'  hi. .oil  oi'  maiix  pioleins  clicmi 
cally  in.lisliii};iiisliahle  from  those  ol'  the  lilooij  seniiii  itself.     Wlnii  jiio- 

ii'iii  is  taken  in  the  usual  ainoiiiils  liy  nioiith.  these  puis. iis  reactions 

ilo  not  siipei'vcne,  even  snake  venom  is  harmless  » lien  swallowed,  nor 
Is  it  possihle  ilnrin.j  iliir.'slion  of  a  prot.'in  ni.'al  to  iletect  food  protein  in 
the  liloo.l  hy  means  of  the  precipitin  reaction,  l-'inally  it  was  tiiscovereil 
that  the  very  >lou  intiavenoiis  injection  of  ci.iiipletel\  (liuested  llesh  did 
not  produce  on  the  part  of  the  liod,\  any  of  the  reaetioiis  that  injected 
protein  its.-lf  pioilllces,  indieatiiii,'  that  perfect  assimilation  had  occurred. 
l-"roiii  tliese  and  similar  oliserv  ations  it  soon  liccamc  clear  that  i)ioteiii 
.•an  not  he  ahsoil.ed  as  siich  from  the  aiiiiientarv  ■•anal,  Imt  must  first  of 
nil  hr  roiiiiili  tl  hi  hniLi  n  ilmni  iiihi  llii  luiiiiin  iiriils.  iclu'rli  iin  llii  ii  nhiiilt 
mid  till  jiriitini  iif  till  iininiiisiii.  'I'll.'  ili''eci  exi.i.'iie.'  for  this  important 
I'liaiijte  ill  helief  eonceiniiiL'  protein  iiictalioiiMii  has  lii'in  '.Mined  by  the 
iliseoverics  that :  i|  nilroircn  .'.piililp  iiim  can  I.e  maintained  in  anin  Is 
fed  with  eompl.'t  ly  diiri'ste.l  pn.ti  in  iiii.xtuies;  and  1 2  amino  aci.ls  ran 
lie  isolated  from  the  hlood. 

The  cxperiineiits  of   the   first    ■_'roiip   .'onsist.    in    principle,    in    lireakinir 
il'.wii   protein   unti!   tliere   is  no   hni'^^-.-r 


I!"    cSui! 


liiUi'i't    te: 


SI,  anil 


lion  feediiii,'  this  disjestion  mixture  to  animals  and  ohserviu..'  thoin  from 
day  to  diiy.  usiii-r  as  criteria  of  their  nutritional  condition  the  hody  woipht 


Mm 


ii 

M 


i     '  ! 


I    '^  ^ 


.V»(i 


Ml  TAIKil.IsM 


^""'  ""    ""'■""'' "'Ii""-ii'in.     il'iii:..  :,7I,,      It  \u,s  „  sl,.m„  il,;„  „i, 

'■"■^'  niai.i.ntf  iM,l,iti,„ml  ..ftl.-i.  ,„.y   .|,.,,..n.|s  |mitlv   ..„    ,1,,.   „at,n 

"'  'I'"  I"-.-  i.snl  r.„  .IkTMi...,'  th..  ,„■..(..,„.  ,.,„|  p„,,|v  .,„  iIh-  ,.r,.s.,„ 

'"■  "'"'"' '■   •■"••'"•Kv'inilr    in    th..   ,lip.,ii.,„    ,niM„n.       It    «as    f ; 

Hiiil  .mmI  I,v,|,„|_vli..  ,„...lu.-t>  l„il...|  t,.  tn.iint.nn  <..|.iilil.i  i,,,,,.  ,,...1  ,i   u„. 

,:^"''  '"'  "'••"   "''^  ""^  ""  '"^'  '"  'I'"   ••"'•1   lIlMl    tl„.  ,,-i.l  h.Hl   MMMV  ,;,mi,\rU\. 
'''-"'""'    ""•    '"•"••■i"    •""l'-"k    ••IM.I    lu,.|    l..tt     .„.    |,n|v,,..,,ti,|....    «l,i,.|,      i, 

""-  ""•'-""••'.  •■'■""li I  intiiH  .luri,,.,-  ..m/.v ..Hi..,,  mi.l  ^yvv,■  ,.ss..„ii,l 

';"■  '•';'•""    '"■"'"'"   ""•••■''•"'!  •"      This  xi..„    hns  ,„.„    I „  ,.„„s^|..r.-,I.K 

i'll''--..!.  s,n,.,.  ,t   l,,,s  l„...„  >h„«n  thi.t   th,.  ..M.i.l  „..t..Ml|v  ,I..slr..vs  vrrUw. 

;"";""  """'^  "''i'-''   " «-0<'>-  l""VH  iMl;,-l.     Th..  „mi„„  ,,.-i.|    ,,;mi,.n 

';"'>  '■'" •■ '  '^  I'M-'-'I'l'-'iMo.     Tims.  ,vh...,  ,„,i,„i,ls  u..,-,.  f..,l  with  Ihn, 

.li.'N.  ..„nM,ti„..,  „(  n<    fully  ,li,-..st...l   ..„s,.i„.    r.>*    fullv   ,li«..s|...]  ,.„...i„ 

Irnm  «h,..h  th..  lrv,,l„ph,m,.  ha.l  I n   ,•..,„., v  ,.,1,  ,„„i  , :{)    fuljv  .liif.sl..! 

'■•''"■'"    '"""    "'•'"•''    ""•    "M'l'-l'li.Mi..    ha. „    ,vnM.v...I    Mii.l    ll„.„    ||„ 

l""l";;'  ■"•""""   '-•■  I'M.-  t,v,.t„|.han..  a.hlr.l.   it    uas  f |   that   .,itr i, 

•'''""" '•'  ""•  '"•  •""i-'l.mM.I  .,„  th..  s n.l  .|i,.|.  whi..| ,lain,..| 

;;,''  ">l-t"l'li.m,..  „|„.r..as  it   was  n.aiiitai.,...]  ..,.  Ih..  first  a„.|  Ihinl  ,|i.t- 

.""'    """  'M'liiMati s  ,....r..,-t    is   fi„-,h..r  s.i|.|,„it...l   l.v   Ihr    fa.-t    tin- 

'I  •'"■  l""l.'ni  .s  .„,ly  partly  ,|i.,r,.st,..|  |,y  ari-l  that  is,  ,„.t  .Ii-,.,,,.! 
•••H.uuh  s.,  as  t..  I,r-ak  .i|.  all  th..  try|,t.,,.han.-      it  ..an  Hll,-i..Mtlv  ...ainla,,, 

IHl?nu-r lUililirilllii. 

Iff-MMliM-  th..  iH.....ssiiy   f„r  ..arlM,hy,Ir,it..s,   it   is  p.,ssil,l,.  that    rni.!.. 

'■"'■'•"' '"I'ti'ms  th.s,.   ,„ay  |„.   pr.,.lu..,..|    fr.„„   th,.   p,,.t..i„   its.lf       \t 

I.^ist.  ,1  h.s  Im..„  p.,.sil,I..  f.,r  .\lHl..rhaM..|i.  wi...  has  .1,.,...  a  lar^,.  shar.- 
"I  tins  M.,rk.  t..  maintain  an  animal  .  nitr..i:..M  ...|.iilil.ri.,m  with  a  -Ii.  I 
"I  .li'-'..sti.,n  |.r,..lii.-is  Mil.)   fat  (-..iitaininir  im  (•iirl...liy.lral... 

Th..s..  r..sMlts  .,l,tain...|   in  .lilT..r,.nt   rlasM.s  ,.f  animals  havo  als.,  1,,.,.,. 

"""",'■ '  '■"•■  ""•  '"'  ""'•'•i'-'      A  l..,y  sn(T..rini:  fr.,m  a  stri.-tnr..  .,f  th, 

.■s.,pha«„s.  uh..n   f,.,|  l.y  r....tnm  f.,r  tifl..,.n  .lays  with  .liirosti-m  pr,..),,,.!. 

*■':""'"-   '■'■'""  ""■  ""'i •'  "vpsin  an.l  rr..psin  „„  ||..sh,  j;av..  .vi.],.,,.. 

',1  riitr.iifi.n  r,'t,.nli,,n. 

*■ '"'"-  ""■  ^"'•■■"•1  'M"-  "f-vi.l.'M.....  many  inv.-st i._^,t.,rs  att.mpt..' 

to  s.'Paraf..  th..  an.in..  a.-i.ls  th..n,s..lv..s  fr„m  th..  I.I 1.  parti,.nlarlv  ,|,„ 

inir  th..  .|,-:..st,.,M  .,f  a  iar-..  an.,.nnt  ..f  |.r,.t..in.  hut  tl...  r..snlts  w.-r,.  m 
(ir.t  .ntmly  n....MtiN..  I.,.,.ans..  .,f  th,.  Ia..k  ..f  mHh.Mls  that  wr,.  s,HV 
'•'.■nily   .1.  l„-at..   t„   n.ak..   it    ,..,ssil.|..   f„   ,1,,,,.,.,    ,i„.   ,,i^,,„    j,,,.,.,,^,^^,   „,.,, 

' '''   '"■  "^■'•■■•"••l   "V"M   «h-n   a   maximum   al.s.-rpti f  nit. ,   h,„! 

'"■'■'"■'■"■'      ''■'"•  '"■>    '•"•'■'"•  """•  "(  I'l I  tln-.n..rh  th..  p.utal  v..in  -aus,- 

su.-h  ..Nt,.ns,v,.  .i,luli.,n  .,f  any  sul.staM,.,.s  a.M...i  t.,  it  that  th ..•••ntr, 

'"'"   "'   "'"  ^"'-''""•<'   "1   "M    is„la....i   sampl..   ..f  th.-   1 1   ,.an    nh 

trivial. 


TIM:    Ml  TMliil.ISM    OK    IIKtII  IV 


'.97 


l> 

f..|||„ 

.' 

■    HIS   iif  i|c|pi';ili' 

'i      y    „ 

IMIMP 

ii 

ii  ill  till'  \<\ 1 

-lill. 

■-.    (   '■( 

II 

I'iiM'cl    ill    the 

1    .11   ' 

■    '    ■  'H 

1.          •IM 

til 
ii '  II 

stilir.      Tlli'V 
il    till'    piirf.il 

'  M. '     ; 

.■1  1-  .■    1 

M 

1     .'iIisipiIh  (I    ,•|^ 

.hi.-^.; 

.'.ill-' 

liv  tlic  iiclimi 

T<.  afi-oiiiit  f.ir  III.,  imlispiitjil.t.'  •liMi|i|M'iiiiiiiri'  ..I'  th.'  iimino  iwi.ls  rrnni 

tin-  intfHiilir  .liiiiiiu'   piMtrili   .liirrsli.  II li|ili'.|    with   thr   ilil|M.ssil)ilil y    iil' 

.l.'fi«.-tiim  any  i,f  iIkhi   mi   tlic   hi 1,  Uv  „  ^,\,.\\H  \\,'y>.  niri'i'iit    Cnr  iiiiiiiy 

vrars.     Iliic  nf  iliis,.  „,.,s  ||,„t   III,'  .iiiiim.  ari.ls  iMi-.iinr  ilraiiiiiiatnl   iN'M. 
i.'r..iip  spill  up  as  Ml    >   |,y  ,|„.  iiii,.sii„,,|  ..pilli.iiiiiii,  aii.l  lli.'  otlM'f.  lluit 

tins.'  .'••IN  aif  i'ikIow,'.!  «  jil,  ||„.  p„u.r  „(  i iisliiirlini.'  tlif  aiiiim.  a<-i.U 

info  prHf.Jii.  «|ii,.||  tlii'ii  passes  int.,  tli.'  |,| |.     .|iisiili..aii,,ii   f,,,-  ||,..  ,|,.. 

aiiiiiiafioii  li.v  pi.tlicsis  s.'.iii...!  |,,  I Iitaiii.'.j  l.y  the  ..l.s.iv  ;ii  i.,ii  iji.it  i1,.t,. 

is  Miorc  fic..  aiiiiiinMia  in  tii.'  Iil.,...!  ..f  ||,.       ..(.i!  v. 'in  lliaii  in  tliat  «(  tin' 

s.vst.'mi.-   .•in-iilati.,ii.      Tli.-    falsity   .i,  ,,.„<■..    u,ih.    Ii.iw.v.t,    .l.'li- 

iiilciy  cslalilisli.Ml  l,\    F..liii  find  lii'i 

ijiiatifilativc  iiii't|i,>,|s  r,,i-  Uif  i  stinii"     u 

tluit  tin-  aiiitiiiiit  .,|"  iicilli.'r  of  |1  ,, 

poffnl  liJiM  (I  .jiiiini/  alisoi'pii.iii  .        •  ii,, 

iiiailc   llif    I'lirlln'r   iitipurlant    •        <  . 

Iiliioil   is  rcillx    M'ly    Mill..   Ill  iiiii. 

Midi  finiii  tin-  inti'stiiial  liiiin  i     •   I  .i,- 

iif  ptiti'fl'acti\.'  Iiai-lcria. 

N'lir  ciiiil.l  any  cv  i.lcn.'.-  li,.  t,|,|„  ,,,,1 
llio  alis.nlic.l  aniinii  a.  Lis  Imi-i.iih'  Ini  ■ 
iiil'i  prntfins,  wlii.-li  jn..  then  Iransl'diin 
This  liyp,.|ii.'sis  was  l.as...|  cntiiclv    ,,n  n.uatlM'  (iii.liin.'s,  ami   hail   liii'iv- 

fnr.'  to  !..•  .Ir.ippr.l  wlicii  .lisc.iNcn    uns  nia.ji-  ,,f  tin-  artnal   pi.'s. f 

.'iiniim  ai-i.l  in   tli.>  |>|i,.i.|. 

This  lirjcl"  hisli.ii.Ml  survey  nf  ihi'  siil,j.-.-t  l.ilnos  ns  t,.  a  pi,sitl.,n  uIumi' 
uf  may  pni.-.M-il  t,i  .IIs.miss  tlie  pics.Mil-.lay  l.'a.liinjr  n-_'at<liii..'  pri.tfin 
liM'taholisMi.  Uri.dy  slalc.j.  this  l.'a.-liili!;  is  In  ih,'  clTcct  that  tlu  /„•„/,,» 
ninliiiili  is  hroLiii  iloiiil  iilln  its  lilllnnil,  hiiihlliK/  shiiKs.  th,  ,i,n!iio  mills. 
hil  till  illi/isliri  iit:i,niis  »f  Ihi  i/nslniiiih  sliinil  l,;iil.  ,iiiil  llnil  tins,  iniiiiii, 
■iilils  iiri  iihsorhni  ml,,  Ihi  hloml.  hfi  iiliiili  Ihi  ii  nn  nirrinl  li,  Ihi  nirioiis 
iinjiui*  anil  tissii.s.  nhirh  sifl  „„l  tin  iiniino  iirlils  iiml  ii^i  thiisi  „f  Ihi  ni. 
irhich  Ihii/  rniiiin  /„;•  Ihi  n  imisl niilim,  of  Ih,  ir  hr„l.,  iiilmn,  ,,i;,l,iii. 
Ih,  ,n,ii,i„  ,i,i,ls  „„l  ,;,iiii,;,l  !„r  Ih,  /.c,,  ,  .o.  „/„„,,  ,rilh  lh„s,  irhiih  mivi 
!',  lilunili.l  in  II,,  lissii,s  Ih,  Itisih;^  Ini  ilisiiih  i/nil  i„,i  ,,f  lissii,  mhins. 
'ire  Ihi  II  spin  iiili,  hvo  iiorliniis.  om   n  jin  si  iiliil  hii  iimmmtiii  mi, I  „lh,r 

li'l  thr  ri  miiiiiili  r  uf  Ihi    iiiiiin,,  miil  nml, ,  uh.      ■Ih,    f<,nii,  r  is  i.i,,,l,,l  us 
nrrii  mill  Ihi   liillir  is  o.ii.li:,  ,1  I,,  j,r,„l,i,,   ,  „,  rini. 


I    I  .i\  pi 


h.\  pi, til.  sis   that 
tiiial   .'pithi'liiiiii 

!\\      ll\      till-    llllllllj. 


CHEMISTRY  OP  PROTEIN 


'l■:^ri^^^    t  lit'    r\  t 


l!.-!uf.'  ;:r;"-iM'.itlijj  t..  MiM-ii>N  I  itt-  .••,  iiii-iii-i-  lij..,|i  \\\,i,'\,  \\u-  jiImim.  ,-,in. 
•Iilsii.iis  ,|,.p,.n,|.  it  uill  I,..  nc,-(s.sary  t,.  .•i.nsiil.T  sen,..  ,,|' the  iih.st  iiii|...r|,inl 
I'.ifts  ciiiu'cinini.'  the  .•lieinistry  ..I"  tin-  pnitciii  iii.ilcciilc.     We  sliali  r,N|iiiic 


I 

1  • 


m 


,^  M 


ll||,^1 


I 


'iil 


l. 

1 

1 

f  ■ 

I .-.-.  i 


lli 


-i^iS 


MITAMOI.ISM 


tills  iiil'.piiii.-ilioii  111. I  (inly  ti.  midfrslaiiil  the  liiston  (if  |ii-otciii  in  tl,. 
iiiiiiri;il  ImmIv,  I. Ill  ;iU,,  tc,  follow  iiitclliL'riitly  tin-  irnporliiiit  work  fli:ii 
ll.-|>  ;iliv,'i(l\  Ihtii  ,|iv,-i|vs,.,|  .•.MiriTiiiliir  til..  r,.|;itivc  N.illlc  <i(  .linV-rn! 
prulciiis  ,is  fn,„l,  A  kiioulfilir,.  ,,|-  iMn|,.ii,  cliniiistry  Ims  <'niuc  tc  !"■ 
I'sscnlijil  ill   piiK-iifjilly  all   iMaiicli.'s  ,,1'  iiii'ilical   si-icncc. 

I'n.t.-iiis,  like  siaivli.s,  aiv  c-oiii|i,.s,.,|  „(  niiin.'rdiis  smaller  Imil. . 

Ill  the  cas,.  of  staivli  tli.sr  iMol,.,.|ilfs  aiv  111.,  various  iii..iiosa.'clinri<i,.s 

i.'iii...isi.  '.I..Ntr..s..  .  I.Milos,.  an. I  -alarlos..;  in   il ,is,.  ,,f  proteins  tin  > 

ar..  til.,  aiiiin..  a.-i.ls.     Tli,.  Invakiiiy  apart  „(  tli,.  links  that  liol.l  tlic  iiiol, 
•  •ul.s  t,,.^,.tlM.r  is  ..!r..,-l..,l  in  |„,il,  ,...|^,.s  |,y   111,.  pr.„.,.ss  of  liy.jrolysis.  .., 

'•■■'"'■''  ' ••'"■"•  "I'  ''"•  '■■•"•I  t'iMt  111.,  ivai'lioii  .■.Misists  In  tin.  taklnir  ii|.  of  , 

iii"i<'<'ulr  of  ual.'ral  .'a.-l,  of  Hi,.  pla..,.s  y.Wr,-  tiio  ..liain  falls  apart.     Tlii^ 

liy.lrolysis  nia.v    1 ir...t,.,!  .iilMr  l,y  tli(.  a.-tioii  of  i..ral  a.-i,|s  or  alK.i 

li.'s.  or  l,y  .•n/.M,i..s.  til..  oi,l>    .liff..r..| in   tli,.  a.-tion   of  tli..s..   r..ai;..nt~ 

'"•'"-   "'■"    '"    ""■    '"'■ ••  '■■■>-■   til..   Invakin-  apart    lak.'s   pla.-o   inor 

I'ss   iiMlis..riminal,ly.    u  li,.,-,  as    i„    ij,,.    |..|tt,.r   it    pro,-,.,.,|s   .- or.linfr  to   . 

'I-'linil..  plan,  m  lij.-li  v;iri..s  s.,ni..ul,at   uill,  ili,.  typ..  of  ..n/y,,,,.  ...nplox...! 

•Inst   as  a   .•li..iiii,.al     kn..ul,..l-..    of    i|,..    strii,.|iir,.  of  siiu..,,-  op   ni .sa, 

(•liari.j.s  is  ill,.  I.asis  .,f  ,-arlioli\  .Irat..  .■li.iiiist  ry.  s.,  is  that   ,.f  the  aiiiim. 
ari.ls   111,.   Iiasis   .if   pr.it.. in    ..In'inistrx . 

Amino  Acids.      Tli..r,.   ar...   s.>    far   as   kn.iun,   ..i..ht,.,.n    .|il1Vr..|il    aiiiin.. 

■"■"'■'  •■'" ''^''1  ill  ""•  '•oiisiitnti r  i.rot..iii,  lint  th..y  aiv  all  alike  i 

thrir  ..hai-a.-t-risti..  st  rii..tnr,..  Tli..  most  strikinir  .•liaia-t..risti,.  .I..p..|i,l 
..II  th..  pr,.s,.|i.-,.  in  tl,..  im.iI....m1..  .,f:  di  an  ami;..,  .,rr,.ii„  with  a  liasii'lt 
'■'""I""-''''"  '"  "'■"   "'■  Jii'iirmnia.  an,!    ^  L>  i    an  a.'i.l   irio,,,',   with   at.  a-'hlif 

'•"'"'""•''''■•  ♦"  "'■•"  "•'  ■• 'i''  i"-i.i.     I.,t  lis  tak..  in  illustration  ono  ..f  ll.. 

s.mph.st    fatty    a.-i.ls      nain.ly.    a....ti...       It    has    the    fornnila    (MfCOOJI 

''''"'*'"""  «'■ is.-all...|  ,„W,o,„/,  ami  ..„  it  .l.-p..,,.]  th.-  a.-i.i  pr..],..,!!. 

"''  " "ip..iin,l.      Tl:..   (11     ._M'oup   is  known   as    m. //,,,/.   an.l   the  -/y-M.- 

firnup  (■SU\   is  atta.-h,.,i  t..  it  in  pla....  ,.f  on,-  ..f  th.-  liv.lrou..,,  ;,toin.<    tliu 
■.'.vinir   the    formula    ( '11  Ml  ( •.  M  )|r.    «hi..h    is   amin..a.-etie    aehl    or   -h 
e.M'oll       If  «..  tak,.  th..  next  hi-h..r  aei.j  .,f  tli..   fatty  aei.j  series,  hnvin  • 
th.'  name  pn.pionie  an.l  th..  f..nii,ila  ('11  ('11  COOll.  its  amino  aei.l    eall. 
iihniine.  has  fh,.  formula  CM  (IIVII.COOII.     .Vow  l..|  us  pla.'e  tlie'r..rm.i 
las  ..f  this,,  tw,.  ari.ls  si.le  l,\    si,!,,  in  the  f,,ll,iu  in._r  „iann.'r: 


NIf      (■    rd'.ii 

I 
.  ;;o"i|'  >      II      (:u-'h]  :;rn,i|, 
Arinii.i.-i,.ti. •;.,;. I 
<  ^Iv. .....ll  ) 


<ll 

Ml,    c    coon 

•:i"  i'i"Ui'.ill.         II       'ti.-l.l -r..ii|,i 
Ariio:..|ii..|,i,.ni,.  •i,j,| 
f:il!inilli.) 


M  ! 


Tin:    MKTAIIOI.ISM    (IK    I'K(»T|:|\ 


590 


It  will  be  oliservcd  thai  tlic  (nily  diflViriKv  bftwcfii  the  two  ncitls  is 
ilt-pciulcnt  ui)()ii  a  cliaii^rr  in  the  ■;riiiii)  that  is  attai-licd  ti)  the  upper  vorti- 
cal valency  lioiul  ot"  the  ctiifra!  eailMin  atom,  which  thcrefdi-e  must  tie 

cniisi(Ieri'<l  as  tl ciitci-  of  the  entire  mcilccnlc     'I'hc  \,iri<«us  amino  acids 

|iio(luccd  fi'oni  protein  ditl'ei-  iVonr  one  another-  solely  with  r'etraid  to  the 
'Iremical  nature  of  the  ^rmrp  that  is  attached  to  this  ver-tieal  valeri<'y 
liorrd.  Kviderrtly,  therr,  llii  niittiuiis  llial  niiiiiin  mnls  /ji/.sm  \.v  in  nmnnon 
must  deperrd  orr  the  terminal  uroiips  coiitairrinu'  the  c-ailioxyl  and  aiitiiio 
lailieles,  wher-eas  tin  ihiiriuli  rhiii  rimliitii  nf  imli  of  llii  lii/lilun  miiiihi 
iicids  must  depend  upon  the  dit1Vren<-es  irr  the  radicles  attached  to  the 
up|)er  vertical  liorrd.     This  may  lie  lepieserrted  thus: 


Aiiv  raiiifle 
I 

I 
u 

Aiiv  iiiniiKi  ;ii'ii| 

The  end  ^'rmips  endow  the  ariiirro  acids  w  ith  the  po\nT  to  <'oiiil>iiM'  v\  illi 
lioth  acids  and  liases.  With  acids  they  iichave  \\U,-  sulistil  iiled  niiiiiioiiias 
to  foi-m  salts,  which  earr  iorri/e  into  the  amino  acid.  ,is  the  catimi.  and  llic 

acid  trroitj).  as  the  anion.     With  liases  tl arhoxyl  i.'ron|i  reacts  to  t'oiiii 

salts,  which  yield  amino  acid  as  the  arrioii.     A  most  impor-tant   i-iai-lion  iim 
sisfs  tir  lilt    ii/iiili  iixiiliiiii  iitdlih  Ill/ill  s  iillli    III!    iiiiiiiiii  i/niKii.      'I'liis  ocriirs 
Iiartictllarly  readily  with   I'oi-iiialdehyde.  water-  liein^r  elimirrateil  in  the  re 
.iction,   and    the   basic    rratirre   of   the   amino   acid    lieiiii;    thiis   deslioved. 
I'poii  this  ri'ac-tion  (leperids  the  method  ot'  .s^iir-eiiscn   fm-  determinin./  lli,' 
iMioiirrt  ot' amirro  acid  in  a   miMirie   i  si  e  paixe  (lirti   .      'riir  titration   is  pir 
lornied  by    renderiiiLr  the  solutiorr    of  ariririo   acids   neutral,   tlien    addin-j 
rorinaldelryde  and  titratinir  with  standardi/ed   aiicl.   irsin.,'   plienolphtha 
liMU  as  the  indicator',  arrd  tlirrs  lirrdirry  to  what  deL'ice  the  ai-idit\-  of  the 

mixture  has  b irrre   increased   as  a   result    of  addiiiu:  the    I'or'm.ldeln  de. 

■•sjiree  this  iner-ease  iir  aciditx'  must  depend  upon  the  niimlM  r  nf  .imino 
-'loups,  it  furnishes  irs  with  an  indirect  estiirrate  of  the  com-ihtiation  n( 
the  uniirro  acids.     The  reaetiorr  is  illusti-ated  by  the  ei|u;rtiort: 


?      11 


ni.li.lc  II  iiiJi,!.' 

I 

Ml      r    ((lull      II     (■      tl  III         N     I-     ririijl      II  (I 

II 


U 

'  .'I  III  I  MO  ;irii|  ' 


l'MinK>M.'l,t.|r 


.Another  reaction  of  airiilio  acid  .if  piiysinlniii,-  in','  (  st  is  lli.il  kirnvv  n 
as  the  fiirhiiniiiiii  niiiiinn.  rorisisiiiiir  in  a  irnioii  of  tli'  amino  arid  wiili 
ealeiiim  and  earbonic  acid      Finally,  it  is  important  to  note  t|i;it  tin'  .niniio 


H' 


1 


i 


1,'i' 


; 


j'l 


^^;-i 


!       I 


» 

j 

1. 

•i 

r 
1 

COO 


Mi:T.\ii(>i.is.\t 


L'lni.i,  ,s  x.n    lirnily  .■,tl,M-l„..l :  it  n..nMi.,s  intii.-t  i„  ,M-i,|s  ,■,.,. I  alk;,!!,..  ;,„.! 
iMv,M,.x.-,l,l,.  ,.nlv  l,»  ,,  |.i.M.,.>s.,r..xid,.t„.n.     Tins, Mil  !.,•  ; ,,lis|„-,|  l.> 

tlV.ltlHL'    III.-   ilMlinu    JM'i.l    will,    .SM,.|,    IVM-rnts    MS    I,  V,|  ,•.,!,.■„    ,„.roxl,|i.   ,„•    ,„.,'• 
MUMLMMMl,'.    Ul,..„    ,1:..   ,„„i„„   ;.,■.,„,,    i.  ,|  is|,ln.v,l    m„l    ,-,    s„  .Mil,.,!    k,.|„„i,.    ;„•,.' 

'"■' '■     ''"'"'   '■•■•"•"""   "ill   I'.'  .vi,!..,,!    from   tl...  i.nM„.,).anv,M^.  .■,,iiiilin„ 

<'ll.  ri, 

"       -"^'l        '■      <'><)ll    .  -•   (I        <■      ((Mill        Ml, 

ir 

T"  illHstr.1,.  this  n.„.|i,.„  u,.  |,;u..  clinscn  .,„inn|,r,.,,i„„i..  ,..i,i  .,,  ,,., 

'"";■' •"'-•!'-"l-'^i '•'"■'"-Il'.v   i'-.N.h.tiMn.n.|k„nu„;,s|,v,,n,',. 

•■"■"     "  "'    "'''>'   -"■'"    ' ■•'■^■•' ill   inl.M„i,.|i.-irv   mi,.|; JiM,,       |t   mt^,  . 

:"  " """""  ^"'-''"'-■-'  '■'■"-1,  ul,i..|,  ,,n.t,.i„s.  ,.,•„•|,„|,^,|,,,l,.s  ..,.  ,-,„s  „,    . 

"   '"'"''■■'■  '■""'   ""■'•'•"'I''  =1-  111.'   iM..-i, |i,,v   sul.sl.,,,...   tl,r„„.M,    «l,i,.| 

'" '.  ""'"  '"•■'>    l'-'^^  ""  '"■'"-  '.■■.■inM--nn,.,|  i,,.,,  mimmImt  .  ,„-,l'c  tidlli       Tl.. 

!'"■.'"   '""  •"■'■""  ^  l-.Mtint:  i„  .., sit..  ,lir..,.ti..Ms  i„  tl„.  :,Ih.v,.  ...mali... 

'";;-'-  'I-.   <l''----.i,,n,n:,y  , .,.,,....,., I,,,,, i|vi„„i,„„,,, i,,..,i„„ 

'  '-;  -'.,„„„„.  -..,  tV,...  IV.,n,  .,„„,..  M,-,.ls  „M,y  I,.  ..Ni,ii„,l  ,„  ,■,,,.  „i„, , 

,';,"^"'"  '"" ^  ■•"■i''  •• •■•'"'-  •"  tl„.  u,.,„.r.-,l  ciuMti,,,,:     Ml    ;  ||(.\o 

\,\       ''";"    "''^   '■'■■"•" '■■'"•"■1^  ••'"""''-•    "Mmn.-lv     i,„,..,>l,„,. 

''"";'.''•"';:'  """"""•'"• ■-///.. /,.,•„/■ ,• ,,,.„„,,,,„ ,  ,„ 

'"■        "      '""   ■'■'"'"    •     ■'■"   ""'l^''   ""■  -ti,n.-,ti.,M.   nitn.us  .Mci.l    is  .•,||„u,.,i 
'"  ■•"■•  ■■"  '!"•  •■"11"-  .i-i-ls.  :,„.|  tl,..  x,.luM,..  „r  nitn.u,.,,  .^^  set   tV..,.  I,v  tl,. 

'■"■"■'. '^  •"••"^'"•,■,1.  tl,..  ,„■ i|.l..  l..,„^.„„il;,r,,.  that  „s...i  f„r,|„.  .1,. 

''■nHUiati-.i,  ..1-  in..;,  I,\    tl,.-  I,y|..,l,n,init..  Mi..||i.,.i. 


Ml.     ,ii:|.:,i;.t 
II V..IU-!    ,iit- 


■  I    f. 


■'■"'l-i,-'  tl„.  M,|,.i;,,„.,.  „„.|  ,-.,ll,.,-,i„^,  :nu]  „„.:,.„,i, 

„, , .,1,  ;:::;;,r;:''^''':;''''''''^'''V''''''' •■■■"•••' ^-'--r 

.1 :,  ■  ;■ ;':■  ■■'"'"■' ^'-'" -■ ^ :■  ..^.i Jl „. 

,      ■  I-    -i'M   •-.   .,. „   M„.l,l,,!,„„.  ,|„.  ,„i,i,„_,  I,,,,,,  .,  ,i,^,  „  , 

'"  ":""    ".:,ik    ..,1  ,    mi|.,i„  ,,\„|,    ,..,.        .    „  ,  .  ,  .     ,  "" 

,,   ,1     ,     .H  I.  ,  ~  "-':i-iM'.l   .|ii:,i,l,n    ,,|    th,.   ;,ii,„  ,   .   |,,i|,,. 

„:,.,■,■      ••'-■""■'-"-"1    -i- ri„,„,|„-,.Tn,:,„.:,„„„ 

,    ''•'"■  ;'l'l-'-'--'-i-"    l-n  s„  ,„.M',.,.,,..|,|,„   nM,„..,..„Kanalvs,.s,„;, 

:::::''■""'"""  ■'^•■'■^^'-'■'" -M.,tl,a,l,..,.,. •a..,.nfa..vtl.,„ 

IS  s,.,„-,.,.|y  s,n-|,..,ss..,|  ,„,,,>   ,.'lu.,-  I„„..l„.,„„.,-,l  est,,,,, -ion 

,i,'7;'' ''''■''■■'''' ■•'::'':^^  ;■'■'''•■■•'-' '•'•--'-. ^t  si.„i,i..aMt , ■.-.,. 

" '""■'■"" "'-^  "'<■"■  ^il'ili'X"  link  „.....,|„.,.,..f„n, , un.l- 


Till;    Ml   I  Mill!  ISM    (iK    {■Kdll  IN 


&n 


.•.illcl    liijtiiil...      TM>    liliMlM,'    nrs    I..  tvM-,ii    lli,.    ;i|ilil:n    -icili)    ,,(■    OIK' 

■  iiiiiio  Jic-iil  ;iii.|  III.'  .•iiilp(i\_vl  u-i.Miii  (if  tl ilicr.     When  iiliiiiiii.'  :iii<|  jilvcu- 

roll,  uitli  wliici,  uc  .-ii-.'  r,iiiiili;if,  ;ii-c  iliiis  liiikrd  t  nuvi  li.T.  llic  rcaclion 
i.ikrs  |)l;icc  ;ifciiriliii'_'  In  llic  ciin.il  inn : 


CII 


lluoc     (•     x 


11     11(1    (> 


11  III  ;i 

'        Ml  11 1 II  M       ( ■      Ml      I  ■(  )     ( •      Ml        1 1  (  , 


il 

(fdlinini' ) 


\ 


II 

f;;lvc. Ill 


li  II 

f..l:ili\l         -U,    i-.ilj 


III  lliis  tiinMiicr,  ilicii.  a  sn  .•allcl  .|l|,c|,n(l,.  Is   ImiimcI.   in   \\\:\,U   it    uill 

!.(•  (iliscucl  tlicrc  still  icinaiii   iicc  .mi-Impnx!  and  ai -rnii|,s,  ||nis  |>cr- 

iinltiii._'  the  liiikin-  nil  „\-  nil,,  •■  aiiiinn  ad. I  -i p^  tn  inrni  1 1  i|,cpt idc-.  nr 

'.  lia|..-|.ti(|,-s  ,,,■  ntlici-  |M.|v|,,  |. tides  IndcMl,  ,|,;s  prnccss  ,,|  cniidcnsa 
■mil  iriay  'z«  nii  prad  icall\    indclinitcl  \ .  a   pnl\  prplidc  cnni  ainiiiLi  ciLdil.M  ii 

'nlill..ilcid  -rnlips       llalnclv^  tin leucine  and  lil'teen  -    ^enenll  ..-rniips       l,a\ 

"'-'  iirtiially  I n  s\  ntliesi/ed.     ■||ie  iv-.iill  int;  pnlv  pep'    |es  lM\e  the  prnper 

tics  of  derived  prnteilis  like  t  lie  pi,,Ic,,.  s;  Inils,  lhc\  -\e  tile  l.iiircl 
-n.j    ntlier    reacti,,iis   (diai-acl    iisiic   ,,|'   proteins   and    are    preepitatc.j    l(> 

-iicli  rcairciits  as  lemie  dik^ride  and  p!i   spl,,  t  ninjsi  ic  acid.     Sniiic  ai" 

lis..  dJL'.'Ntcd  l.y  trvpsiii  and  .repsni.    Tl  e\    l,a\e  thcsa ..ptieal  aclivitie- 

-  prctcins.  (Inc  ..f  ilnm  I, as  l,c(  n  pi,  pale. I  w  Meli  pr..diic.-s  a  typiea' 
.niapli\  lactic  rcaclnni.  -,,  la  r  a  p..l\  p.'pl  iiie  that  can  l.c  cna'.^iilatc.l  1.) 
!.;it  nr  is  in  nil, cr  \>a,\s  id.'ntical  \>itli  the  natiirallv  ...-.Mirrin.j  pr(.|eiiis 
'.!>  lint   lien  sMithc   i/ed;  l.n'    tlicrc  is  i„,  d..iil.t    lliat    it    is  nnlv    a    mailer 

■l'   time   hefnre    tilis    \vill    I...   ac.-nl||  pi  isli.^d 

KiLditccn  distinctly  diir.iciit   aiiiiii..  acnls  lia\c  liceii   is,,!atc(l   I'-.. in   pr.i- 
'.■III.  and  it  may  assist   in  tl  ,.  (-..nccpt  i,,,,  ,,i'  .mr  prdil. m   it'  w  c  place  tlicse 

iniiiin  a. -ids  si.jc  I,,  side  an. I  lii.l,  il.em  i,e;ci|  ,  ;■  i„  rl.c  manner  d,  sci  ii.e.  I 
-l.nvc.  This  is  ,|,,i  ,.  in  tl...  ,•.•,■, ,, lip  •■  ni-  (diarl  c..n,piie,|  l.v  \)  |i  Van 
^lykc,  III  uhica  also  \ariniis  ,,llic.-  iiiiii,.rtaiit    fads  .•nreernin.j-  tin-  (dieiii 

■try  (if  till'  aliiiiin  acids  ar.'  inci(i.iita  11  \    a.ldcl. 
At  the  Idwer  part  ..f  .'a.-li    f.iiiiiida   \'-i\\   lie  sc,.|i   t|..'  elia  ladeiisi  ic  car- 

■■'\\l  and   aniiii.i  .ji..iips  ..f  nciLdil...rii!i.'   acids   liiH  iin,'   Pejcl  her   ihe   icr 

■Miiial  carl.nii  at..iiis.    'Id  e  upp,^,-  \.'rtl.-at   I  1  ,.f  ||,>s  earl t.,ii,    is  .-.m 

':..-tcd  with  llic  .■haraci.risii,-  emup  ,.:'  jf.  aiiiin..  a.-i.;.  uind,  ||a.^  l.c  \.-r\ 
-iNplcamI  repres,.nl.'.|  ..nl^  \,\  h  v  .lr...j,ni.  as  in  ulv'"-.'n,  -'r  Vr'zMv  c.mi 
liii'X  and  ilndildinu-  a   rnnj  l'..rniat  i,.ii,  as   in  tivp:    phaiic       It    will    fiirtli.r 

I-  nliscrvcd   lliat    //../■-    ,r,fi  h,    „//,,,•  ,,,„,./..  .//„„/, V  incdcd   in    xaridi.s 

i'.'silinns  in  this  radicl.'      This  is  parli.-iilai  \\   tl ase  in  the  liist  tin i 

■'•'  ■•"'lili'i  .-I'-ids  ill  the  laMc  naiiicly,  (he  l.asi,-  amin,,  acids.  In  i\sii|. 
liic  I'Xtra  aiiiiiiii  u't'niiii  reacts  with  nitmiis  .ici,!.  lilnrai  inu'  fr.'c  nil  r,. "en 


y 


1;:  ij 


!" 


ft 


■ii-        I 


n  . 


mi;tahoi,is>[ 


i    ! 


M!    ^ 

"'lis 


i-q^li. 


;     r        X 


i  a 

,    t. 


rt 


I 

1         ll 

-a  f     -    •.    V* 


=  -^'    5 


- 

|S1 

1 

;,                            t 

3:1? 

i  ■''         ' 

i    '           k     ' 

.»;.  -         ,-— K  •  ■  ■      . 

I   ■.       1    »    i 

;_-/'.'     K* 

'  i  i  ' 

-=-          0      

M 

1    f 

_,,?-S^ =_?_J 

C 

i~i''                        ;           H 

-      .                            = 

£ 

-'':::^-H-'li  • 

-    =                ;       ■■" 

-      ■» 

r              ■  • 

;] 

f'     =  ?    i 

1  i?i-- 

J|;?' 

:     "    . 

rJ=  '  .  .  .    =  ?    .1  I 


THK    METABOLISM    OF   I'ROTi:i\ 


603 


>\  the  Vail  Sl.vke  inftlKMl;  Imt  in  nthcr 


cases,  us  in  aiKininc,  it  fails  to 


till' 


■_'iv(>  tliis  and  tlio  other  t-liaracteristie  reaetioiis  of  tlio  amino  jrrotip.     That 

extra  amino  jfi'oiip  in  lysine  reacts  ilirccily  witii  nitrons  aciil  explains 

why  various  proteins  when  examined  for  amino  nitrou'cn  yield  an  amount 

ihat  is  equal  to  half  of  the  lysine  iiitrotren. 

It  will  further  lie  oliscrvi'd  that  the  iiniiiio  acids  are  arranged  in  three 


inani  iiniups:  one  liasic.  ;inother  ncut 


i-^il     .nul   tlic   third   :o-id.      Tin    mills 


./  till  hiiMr  iinnii,  jirc  three  in  nuiiilnT  jiid  li:i\c  .m  iilkiilinily  siniil.ir  to 
Oiat  of  ammonia.  They  have  Itcen  called  the  /-«,«»<  husis,  hecause  ea** 
'  ■iitains  six  carlton  atoms.  They  .n-f  iilonc  prcsiMii  in  certain  forms  of  pro- 
I'in  called  protamines.     '1  ti>   ntiilrnl  (iniino  iwiih  contain  one  amino  •;roiip 

lid  one  earlioxyl  tjroup.  wliicli  c\actly  n.-utraiize  each  other.  This  is 
iIm'  larjrest  ^'roup  of  amino  acids,  aiiil   is   further  sulidivided   into  three: 

iiic  containing;  aromatic  or  lien/.ene  rin^s  and  includinir  the  very  im- 
|"irtant  amino  acids,  tyiosine  an<l  tiyptophaiic:  another  coMtainin<r  the 
Mi-calle<l  jiyrrolidiin-  rinir;  ami  the  ihird.  the  larjrest  of  all.  containiiii: 
ilie  soealled    aliphatic  chains;   that    is,    the   chains   chaiact.'iistic   of  the 

latty  acids  an<l  which  may  I iih.-r  stiais:ht  or  liraiiched.    When  the  .-hains 

irc    liranched,    tin-   siil.stai is    called    all    isoiilistancc,    as    In    isoh'iicinc. 

Ilii  mill  iimiiiii  Ill-ill-:,  includinir  irliitamic  acid  and  aspartic  acid,  are 
'  liaraeteiized  l>y  containing:  two  carhoxyl  yroiips  and  oulv  one  amino 
JioUp.     They  therefoic  resenilile  acetic  ac'il  in  ac-i(lit\-. 

It    may   he   of  assistance   to   siune    if   wc    restate    these   cheniii-al    facts 
Mom  a  slightly  dilTcrent  stand)>oint  as  follows: 

'''.'"■""■  "I  Jy "II,  is  :itniiii.;i.rlii-  :iri.l,  ('II  .\  1 1  (  1  )l  »||. 

Mt 
■"""""     "^    t:'-*'- V'""    "    ""■'livl    un,!.,.,    rilrir  .    j,    ;,   ,|„.,,.,-,„.,,   ,„„i„„. 


I'OOII 


"pi.iiiii-  !iri,|  aii.l   Is  ilnsi-lv  iiliilr.!  t.i  lii.-tii-  :i.i.l,  \\l,i.li   i«  Cllf'll 


OH 


\ 


.1/100/  I'f 


rooii 

"('■  iitliir  iimini)  ,i,i</.«  imiii  h,   i-,.,i,vi</i  r,,/  „.,  ,i,  riritin.s  ,,f  oliuiin, .  tliii^i: 

1.  Siriho  is  iiliiiiini-  Hitli  :oi  ••Oil"    1  li_vili..v\  1  ,    ;;nmi.  in   |'lii.  ,•  of  uik'  ,,t'   lln>  •■ir 

MI 

/ 

t.im.s  .if  111,.  in,.tli\|   yiHiij.,  ('II  Oil      (11 

\ 
VooH 
J.  ('tjstiiHi'  i.s  alanine  uilli  jiii  ••.SH"    ,  tliinl    crini|i  in  tlii--  |iusitiiiii, 


Ml, 


ciisii    rii 


/ 


coo  1 1 


i!!         i' 


iP        *  .  i  . 


liOl 


Ml  TAIIIIMSM 

T»,.  ,,-t,.i,„.  K„.|,,,.h,  „,„(,, I  M   ,1,,.  -..s-'  _,,„„,.  ..;„.  ,,,.,„„ 
Ml 
Ills     (II 


II  "III 


/■//,  I, III. il, I, f II,,     li:i,   :,   (•  II 


(■h.i.vl  ,    u'..!,..,   CM  (•  II       I'll 


Ml 

cooil 
Ml. 


I.     /■'<!..,.,.,      !,■,,    ..,    (■    II   1,11         |,|„.„,,|  . 


■l'.   >'V.   I-  II  nil       I'll 


\ 


COOII 


/    ir''i'i'"'"  ii'-<-  II  '',11 


•     II      I   HI'l..!.' 


Ml 


•'        <ll        'II       Ml        (IM.II. 


'•II  III 

\ 
Ml 

>ll 

\ 
N  Ml 

''""    t'l-  '•'  ''11  '■  i 1:,/,  I. 

'11 


-n..i|. 


\  Ml 

I  I 

•'M  '•  '11     .    '11      Ml       I  doll. 


IS    illlol 


•'^'    '  "■•'   ■''■^"   '•••'ll'-'i    l"l'r..rii,l„     r,.,i,j„„n„l,.   ,,|'   u|,i,.l|    ti,, 

-.   i'     l"""-  "I'l'li    I-  a   |.M>..|;,|,,,,.   ,•;,:!.,. \,|j,.   :,,„l; 

'■II  <ll 


ml,,    ,        .( ',    ,,/,       ;,, 


I         \::U'r 

I-.i..  , 
1 1...-.' 


Ill 


•  11           1 

lll'lllljl 

.Ml 

I'll 

'11             in 

'11                       1 

II 

III 

III 

\ 
<ll 

1   1!   Ml 

III 

'   11  ,M 

II  Mill 

'II  Ml 

'1)1  til 

v:Jr„'i 

CUUIl 

(U,.|,i„i„. 

) 

(  II  llt'ilh') 

nil      Ml  TMlul.ISM    UK    fiioTI  l\ 


I  L'  I     Till'  IIIIIIIIM  illli.isir   Ml  |>I.~ 


.\>|>Illli.-,    ul.i.li 


I-    ill;:IP|i   >ili,  iliii     :ii  i,|. 


CM   ('(Mill 


Wt.'l 


<ilnl 


III  Ml  rill  III  ;  I 


ittiitiii-,   \v  111!  h 
<ll 


llliilti'-lllfitl  u-     :iri( 


<'ll     I -I  II  III 

IIIMI     IIIIHI. 

I.nxtlv   llM.r  :,M-  111,.  ,li:niii ,.  i,N.  i„  ,,|,„  |,  ,„.,  .^,,.ii|  >  ,.\l.|  : 

l.ii'in:    a  I  ■li:iiiiiii.'.:i|iriih'  :iiiil. 

MI'ii       I'll       III       I'll       ,11      ,.||- 

<  I II  III. 

ti'innii,    n  Mioinii      .1  ^'Kihi.liiH'  \:ilri  ill  ii-   ,•  .  il. 

Ml. 
/ 
IIN-       <•  Ml 

Ninii     I'll     I'll     I'li^ 

I  I  II  I  I  I 

'I'll.'    -liMlii.Jili,'    '.^i..n|.    i:,    fi  I-    :m  I  i     ,.    ..(    i,,t,)i-t    1,.:,    •     ,u    ,] 

Ml 
'  :i,    Hliiili    Nil        I  • 


n  l;tli"ii^l,;|,    I, 


Ml. 


'  {} 


if 


k 


111. 

■J-  J  r  ' 


riiAiTKi,'  I. Win 

Tin:    MKTAhdI.IsM    n\-   n{(»T|.;|\    (CcntMi 
AMINO  ACIDS  IN  THE  BLOOD  AND  TISSUES 

In  the  Blood.     I'iii„i.|,r,|  uiil,  ti,,.  ..-.■ii.thI  f!u-\s  .•... riiiiiir  il,,.  rh,;,. 

I-MV    „|-    |,n,|ri„s,     u,.    „,„y     n,,«      |>n„-r,.,|     to    runsi,!,.,-    th,'    III..!-..    piTci. 

' "''•'''-'<■   >■ iillv    Mc,|iiiiv.|    ,-,.ii..,.|iiiim    til..    liist..ry    of  tliis    siil.^t.' ■ 

'"    ''"■   ••'"'"'■■''    '•'"""".>.       .\ltlinu...h    1 „-•    llil^    sl|,-,M.,.,|..,|    i„    s..pai;,li.  .- 

■■'""""  •"■'''■^  '"  I""- iMlition  fr,,i„  ,|iMUM  I.IcmhI  ..vni  .Iiirinsr  tli.-  lin-i  ■ 

"I'    -li'-'.-tioU.    it    I, MS    l...V,.lH|..|...s    l„.,.li    |m,sm1,|,.    1„    ,|„    s„    IVOIM    ..irriihiti;   - 

l'l'""l  I'V  M  „„.tl,.„|  ,,r  .|i;,l>sis.  ;<n,,UM  MS  r;rl,l,ll,ision.  .■In|„.ratr,|  I. 
■^'"  '  ■""'  ^"-  I'lipilv  TUr  nirtliucl  .•misisis  ill  roniUM-finsr  j,  I,,i,;-  tii!-- 
;'l'-"ll'"l""i  ^^t'li  tl.r  tuu,.n,K,,f..,  ,.,11  ,.,it,.r.v  ill  Mil  an..slli,.li/,..|  amii, 

'''"■  '"'"'■  '■"'''■''   '"■■'">    '"'"•s.   is  III.. II    ii,iiM,.|.v...|   in  M  s.,|,iti..i iitMin,. 

.■M.|MnNi,„Ml..|N    th,.  .am,,  salt   ,t,.|it  as  l|„.  I.|„„.|  pjasina  ..f  tli.-  aiiiii. 

''■'"•  ;l"'l''i-il'l''   ■•nnstiiii,.,,!,  ,,|-  111,    1,1 1    ,,1;,,,,,,,   ,|i„|vz,.   int.,   th..   sal 

""'''"'""  ■■  '"■  ""'   I'  'I"'' ■■<>    !'«■  |'rv\..|it...l  fr,,tii  .Iialy/iii._r  l,v    a, hi 

"'•"   ■'■"•'i'-"l.n'  sul.-t; I,,  ,1,,.  salin..  in  su.-l,  ,.ini,„M.ts  ."is  u  ill  ',„aK,. 

'•■'n,-..|ilrati„ii  in  |,lasi„a  an, I  salin..  alik...     hi  s.mi,.  u;,ys.  it   „i||   |„.  .,., 

""'   ■'''''•■"'•'"I-   '"■■i:     I nsi,|,|.,.,|   as   an   artili.-ial    l<i.lii..y.      Its    ],„•<-.]■ 

'■Iiiii.-;il  a|i|.li,.ali,,i.  f,,i  ll„.  |,nr|„,M.  „(  |...|i|..\  iiii;  |„,is,.ns  fnnii  tli.'  li|. 
is  11,1,1,.,.  iiiv..stit;ati,,n  It  li.is  I,...., I  p,,ssil,|,.  in  t|,is  ua>  I,,  is„lat..  m.\,-. 
of  til.,  aiiiiii..  a.-i.ls  an, I  ,.th..,.  aiii„i,„iia  yi..|.|i,ii,'  siil,stai,.-,.s  IV,,,,,  |,|,„ 
Thus,  alaniii..  .in,!  \aliii,.  ha\..  l»..ii  .,l,taiii...|  as  .•lystallni..  salts.  ;, 
hiMi.liii.'  an.l  ..ivaiin..  .  s,,-  p,.,^:,.  trS^  .  sli„uii  t,,  l„.  pr',.s,.„t  |,v  tl,..|,."iv 
ti.itis.  All  ..I'  th..  ami, I,.  si,l,stMn,...s.  |i..w,.v,m-.  .h.  ii.,f  .lialy/r.  an.l  tit. 
.•N.-..pti.i,is  a,-.,  fiirth,.,.  ,.l,a|.a..t,.ri/.Ml  hy  th,.  fa,.t  ,lial  tli..y  .1.,  „.,(  r.M.I 
■-'IV  up  th-'ii'  aii,„i,„iia  i,n  th,.  a,l.liti,,n  ,,r  s,mIiu,i,  .arhunat.'.  as  ih,  • 
-lilViisihl,.  sul.sta,i,.,.s    .I{,,h,l,.   .      AltlM.UL'h   ami,,.,  a, -Ms  .-an   tliiis   l,.<  s,  , 

'■"'"'    '"    ■'    I"""'    ^'■''•'    '■'■"•"    '■iivulatiiii:    1,1 1.    ,|„.„-    ,.,„u...,itfati.m    in 

.Irawu  sp,.,.i,„.,n  ,.  ,„„  |„„   , .,,,„  ,,|,..,.,  ,,„,„„,,„, ^,.  „,,j,„,„|„„   ,,„^^,|, 

\h    III-   n,..||„„is  „,•   Van   Sl>k,.  an,l   S,,|.,.ns..„.   aliva.ly   .|..s..,.ih,.,l.   h.m.-v. 

it   has  l„...u  sh,,\wi  a„i..,iu'  ,,tli..,-  thiii-s  that   th.'  hi 1  ahviiys  cniitain^ 

.■.Tfaiii  .•..n<.,.|itiali< f  ami,,.,  a.'hls:  thus,  ii,  that  .if  fasfinj;  animals  fr 

■'  '"  •''  •"-•   I'"'-   "" •  "I'  I'l 1   lit-'  usually   f.,uii.l   pivs..,it.     Duriiic  t: 

:ihs.,rpii.)„  ..j-  a   [u-..t..|ii   ui.'al.  tl-  amiti.i  ...uit.-tit  ..f  the  Mood  undcrt:.. 


Till      Ml   IMuil  l>M    iiK    PUiili  |\ 


COT 


.1   iiiark.'.l    inciiMs...   1 ninj,   ,|,miI,|ii|    ,tr ;„„]    m    similar   iTMilt    has 

' "    "'''iHIP.il    l.\      |ila.ill-    |.||IV    ;i||llllo    a.   I, Is    III    1|„-    sliiiill     llit.stl.lr.        AfllT 

I'l  ^.TaliiN  nT  Mlaiiiiir.    for  (Aaiiipl.-.    lli,.   aiiiiiio   iiilri...',ii    .,f  thr    iM.'M'iilcri.' 
MiMiil    rose    I'luiii   :!  7   111  ti,:!    nii;     jhi    c',  lit  • 

In  the  Tissues.  AII.m-  rni,-,iim  Hh'  .-ilviilali..,,.  nin..  ai-l.!  r,  ,// 
.,ni,'khi  il,y,ii,i„„r  I'll. Ill  It  a'jaiii  Tlii-  lia-  Imth  .!■  ■  Mialc'l  liv  uIp- 
-rvinir    iIm     aiiiiniiit    nf   amino   aniK     in     \\u-     hid.i.l     .iri,.|-     intrav.' isK 

ii.irctiiiL'  a    ~..liiliim   nf  amiiKi  .ni.l    int..   an    am  stl;.)  i/,.i|    animal       Afltr 
.MJcrtinu:   )•_'  -III,  ipT  nlaiiiiM'  inln  llir  V.  in  ,.r  ,1  ilou.  !HI  |..|-  cml    uas  foMml 


II 


'■H 


ijili' 


I'll;     !-■,      \n,a 


.1    I     I      \l  ,  I 


■■  liav.'  (Iisa|.|..'aiv.|  lV,,iii  lli,.  riivnIati.Mi  uitliiii  li\..  miniil.s      Tlir  ,\u,",- 

""   '•^'   ^^''■■1'    ' 'I'l'-'  "I'  'li''  aiiiin.p  a.-i.N  Iliat    iapi.|l>    i|Ka|i|.rai- ;      Aio 

"■>    ■■'•'•"iii|"'-'"l    in    111.'    I.I I.   .1    ,!,,   tlh'v    l.,-,-Mm,.   alpsoil..',!    \,y    tj,,.   tjs- 

IV."     Tliis  |.r.,l,|,.m  liav  l,.rii  aliack.-.|  l.v   analx/inir  |.niti..ns  „(  vaiinus 

••Lrans  ami  ti-Mics  ivmovr.l  l.rf.ii-,.  ,-,ml  snni.^  tin..-  aft.T  ll,.'  iii.i,...ti.)n 
'"  ""   iiiiiiiiai   (pf  aaiim,   a.-id    sppluti.piis.      In   the   i-a-c  .,i'  the    mns,-l,.s    it 

av  hccri  tiiiiini  that  the  amimp  ai'i.l  .•.pnti^ni  incr.'aM's  until  fi-imi  tiO  tip 
-0    MIL'.    |).-r    cfiT    ,,f    amimp    a-'i.l    has    a.'i-iimiilat.Ml        itryppmi    this    point. 

■pw.'v.-r.  TM.'  mus,.|..s  ,|„  not  s,.,.,ii  t.p  l...  ahl,.  t,.  takr  up  any  im.i-..  Minin.p 

'•1(1.     Thi-  faivap-iiv    .p|'  tin-  iiitcsiinal   .ir>.'aiis,   li.pu  ..\  ,■!■,   is   hhpic  clastic; 

■I  111-    I-    ..    .     iiv.-inrnl    U.I-.     ..f    .I.llns    1-11     lull    ,    ,■     „i    h! I. 


i      1 


M 


In      1 

•I  ' 


' ' . 


H 


■i 


t 


! 
'  f 


1 

1         ( 


fiOH 


Ml  I  \i;iii  i>\i 


/'"■   '■"^■"lll'l'  ■    ""■    iilllil Illno,.,,    ,,f    III,.    lu.T    lljN    l..-r||    ,,l.s.T\...|    I,,    !„., 

iii-M,iM..|  tr.   !•_•:.  ..!•  i:.0  III-    |„.r  .•.■111  ,.|-  ||„.  ,,iimn,,!  ;,iih.,iiil.     .\hli...i,.i, 

tllis     „I,S,„|,,|,,„     ,,|-     ,.,„|i„„     ;„     „|^     I,,       ,1,.,     ,i^^,„,^      1^     ..Ml.||„.|V      .iipi.l.      It      M.     V.I 

l.lM.-.T.Is    I,,    sllrli    ;,    |,.,i,,t     II, Ml     lllr     M j     I  .,,•,, I,,,.,    ..Miilvl;       |V. "    tli,  M 

Mn.m  Jill,  I    in.ni.v    -l.iNs'  stjiiMitiol,   ||„.  |,|„...|  .•nntaiiis   il>     i..n„.,|  ,|,i,,t.i 
"'■  i>"Mi  :l  1,1  Id  i,|....  !,■■  in  I  ..r,||,  ,,|'  „„,■„,  ,j,„„.  ,  |,.,.,    m>^|_     .,.,^1^  in.li,..,.,, 

Ill.ll    .1    r,rl,,in    .■,|l|||il,|i|„||    Mlllsl    |.,V,,||M     .■sliihlisl,,.,!    |„.|W,.,.,I    Ih,     .MIliMn.,      .1 

'■'"'I'-"'  "I'  III'-  I'l"  'I  -'ul  III,, I  ,.\  ll„.  li.>ii,,.  il ,.,-,ilnil„,„  JM  til,.  i,...|.  , 

l'-''N-  .ipin-,  \iiiial,  In    r,,,,„   liv,.  t,,  1,.,,  liiii,.^  iriViit.T  li    11,   III  ti.,.  I.j I. 


'•■■■I     ■!'  I"  I'   \>    \.,„  Ma.  . 


''"'"■     ■''■-■''"■■I     .'"HIH,     .•„.i,U     ;,,,.     ,,.,v      I,,,,.,. I  ,.,,,||l,i|„.,|      „ilj,      ,1„.      ,i.,;, 

>'"■  "->    '-.n,    I,.  ,.Mi-,„.,,.,|    l,v  M,.-li    r,.,.l.l,.   ,,,,,,, ,K  ..    ^,Mi..r  ,„.  .1,1,,.,. 

'■'■'"•'       ''■'""■    l"-^'' '-■>"    '-'      I'"       v,.r,    l„.    ,,K     ,|,„.    t„    ,|i,r,|.: 

'"'■    "    "    ''■■"'     'I n,-..|,;.     l„,i.    ,. I    ,„,,    I,,.,,,,,,,,,    ,_,,,.,■„,.,.    i„    III,. 

^'"7   '''•"'    '"   ""■   '''""I-      'I'''     "I'll..',-   l;il,.  ,.■■   til,.  ...ini.i  .   „,.i,|s   is  .lilti, 

' ""•'■       ^^'''  '■"""   "'•■''  '!"■>  'I"  11.. I  iv„i;i,i,  in  il„.  I,.., I      f,„.  .,  !,„,,_,  I,, 

'"■'■•'"'^"    """•'    "'■   ''"•    l'i"I'iii    hilr,,.j,.,i    ill    III,.    ( I    1.    ,.v,.,-,.t,.,l    ,.|s   „. 

"'"''"   '"■■'".^    '"'"■  ' '-^  ••"■•'•I'  inu.M,..,,:  .-i,,,!   „|„.„   s,,|...l,.  ,,i„i„,.  ;„. 

■I'-'    li'l.  iIm.n    .|I,i,.|J\    i,..i|.|„..,,.  in  til,,  |||. ,„  ,1,.,,,, 


Tin;    Ml.TAIlltl.ISM    OK    IHllTKIN 


(iO!) 


Thf    fisMi.'.s    <i.M    Ihrnlnr.'    I ily    a    slo|.|>iiin-|.laf.'    Ii.r    the    niiiiiio 

aci.ls.  Wh.ii  the  lattrr  iin-  •li'trriiiitM-<l  m  I.I.hmI  i-oll.vt.-.l  fiom  (liff.Tfiit 
parts  Mhilo  alisorplioii  i>|  piut.-in  froni  uii-  inlextint?  is  in  proiess,  it 
Ims  Ih'.-m  I'minil.  as  sli,.«  n  in  I'in  !>■>,  thai  .liirinK  th.-  p«ssatf<.  of 'thi- 
hl.)...l  thr.Hi^'h  the  liviT  th.'rv  is  a  irivai.T  fall  in  tlio  n.nc.-ntrHtion  of 
amino  m.-kIm  tliaii  •InrinK  its  passaj;,-  thiouuh  thi-  entile  n  iniiin.lor  of 
the  IhxIx  . 

It  will   lie  sr.ii   iliat   the  al.ove  .•oiiriiisioiis  aiT  ilrnwn    from   estima- 
tions iiinile  on  l>loo<l  taken  In.ni  ilie  veiin  eavn,  portal  vein,  and  hepatic 


I  i«      IK"       lur> 


^Ii  'WillK    the 


riilt:ili..„    ,,f    .mill,.,  jii.l    ml., .urn    III    llir    l.l.„„l    .luring    fasliiif 
•■111. I  |.r.. 1.111  .liK.,li..i.,     (Fri.m  |>,   I).  \;,„  sivk.-.l 


artery,  tlie  upper  eiirves  in  tli,.  rjiait  lieiii-.r  fn,,,,  animals  (liirinn  diirestion 
and  the  lower  fnun  fastiiit,'  animals.  The  re.sults  sliow  that  the  liver  must 
Im'  parlieiilariv  u'reedy  of  amino  aeids,  which,  however,  must  nipidly  he- 
eome  transfonii.d  int.,  oth,.r  suhstjiiiees.  sinee  no  eonspieuous  varia- 
tion has  heen  found  to  o.-eiir  in  the  amino-aeid  content  of  the  tissues 
aeeordintf  to  whether  the  animal  is  fasting  or  is  discstinp  protein  food. 
This  result,  it  is  to  he  noted,  is  .piite  different  from  that  which  is  ob- 
tained after  the  intravenous  injection  of  amino  ai-ids,  and  the  results  of 


R 

ii!  (1 

,1 
it 

li 

') 

:1      I 

!' 

i  /.? 

I 


MICROCOPY    DESOIUTION   TEST   CHART 

ANSI  and  ISO  TEST  CHART  No    2' 


1^0     If  I-  IIIM 


l.i 


1.25   iu 


>-  Im     ill  2.2 
t    1^     III  2.0 

mil  1.8 


1.6 


A     APPLIED  INA^IGE     Inc 


{7lt>i    288      f)>*8»       Fa. 


hi  ifc,^ 


if^  n 


-8l>i 


i;'':'1 


;* 


III 


Ifi 


!    I 


lllil 


r.ii) 


Nfl   TAIKII.ISM 


ili('  twi)  t'xiii'liiiu'lits  t.ikcii  touctlicr.  imlic'itc  tlmt  tlic  iiiiiiiio  iicids  iil't' 
llii'ii'  ;iliMii|ili(iii  can   iml   iciiKiiii   in   tlic  tissiii's  in  ;i    I'l-i'i.  i'i>iii|iti<iii    (,i\-  ,, 
Inner  tiint'.     It  iiicjins  ili;it  tin   nuiiii,)  nriils  (Jiiriiiii  imlnnil  ilii/i stinn  ;/;..■ 
/)(   ilispiisi  il  (if  (it  (I  r(il(    ii-li((l(  IS  jinicliciilhi  11(1   siiiiii   (IS  IIkiI  nl  irlildi  ,ii 
siivjit Hill    IS  jii'dii  ( (liin/. 

THE  PATE  OF  THE  AMINO  ACIDS 

To  I'.illdW    ilic  iiict.iliiilisin  111'  tiif  iiiiiiiiii  iiciils   furtlHT  we  must  ijct.'. 
'iiiiii'    till'   (ikI    pi-.mIiicI    inlii    wliirli    tlicy    ai-f    ('(HiviTtid.      Tliis    is   urea. 
wliiiM'  cvliiiialinn   can   nnwadaxs  lie  made  \\illi   cnnsiilfi  aide  ii('cill-ac\    m 

■"•••"I"!'   "I'  111'    <liM'(i\,r\,  l,_v   .Mai^liali.  i,f  the  a<'ti<>n  ol'  iii-casc  ii 

MTtiiMj-  its  nitiniirii  iiitd  ; iiinia.  w  luidi  i-an  then  lie  cstiiiiati'd  liy  cnni 

liaiati\fl\    siniplc'   irictlmdN   il-'nlirii. 

W  lien  tilt'  visi'cia  ai-c  cninpaii'd  lii'fciic  and  at  xafioiis  itcriods  at'tc 
tlic  iiitravciiiuis  injci-tion  of  /.nino  acids,  tlic  iiiiiiicdialc  increase  ih 
amino  inti-o-icn  remains  iiniliiiiinis|ie<|  in  ail  of  tlieiii  except  the  liver.  \i' 
^^llicll  a  very  rapid  nchiclion  is  oliserved  to  occnr.  At  the  same  tiin. 
til.'  |)ercenta-e  of  urea  in  tlie  lil.^.d  stc;idily  rises.  Tiiese  facts  are  illu^ 
trated  in  I'iir.   1^7. 

The  simplest  intiM-pretatioM  of  these  resnlts  is  that  the  liver  conveit- 
the  amino  aci.js  into  urea  an.l  dis<diai-es  this  urea  into  the  Idood.  Tlii- 
conclusion,  however,  il  nnist  he  ohservi'd.  is  not  incvitalile;  for  it  is  p.-v 
sihle  th;it  the  amino  acids  may  he  condensed  into  polypeptides  in  tlh 
liver,  jnst  as  snirar  is  condensed  l,y  this  oryan  into  ulycojreii,  and  tli.i' 
the    iiicrc,-|-e    in    ll|-e,-|    is    Irielvly    coincidi'Ut     I  l'"isl;e ) . 

It  must  not  he  ima'.diicd  that  th mvcision  of  tin-  amino  ncicjs  ini' 

ur<'a  is  cM'Insively  a  fum-tion  of  the  liver.  ()ii  the  contrary,  it  is  wil 
known  that  this  proc.'ss  may  occur  in  anim.-ds  from  which  the  liver  h.:- 
Ipi'cii  entirely  removed.  It  is  proliahly  safe  to  concl\ide.  however,  tli.i' 
the  liver  is  the  most  active  center  for  amino  aci>l  transformation  an'! 
uica  formation. 

When   urea    is  estimated   in   samples  of  hhiod   renn>\eil   .-it    sIkmM    inter 

v.-ds  of  time  ;iftcr  the  int'estion  of  a  laroc  amount  of  protein,  if   is  f e! 

that  the  iih-rease  lii'comes  ver>  (arly  eslahlished.  Iu  one  exiieriiiicn' 
liefore  the  food  was  taken  the  I'oncrntration  id'  urea  ullro<ren  iu  the  hliM„i 
\v,-is  a  little  oxer  III  iirj-.  |>er  cent:  one  lionr  after  takiuir  -Mtfl  <ri'!niis  .■' 
meal,  it  had  risen  to  ;ihout  IS,  and  iu  two  hours  to  ncarlv  2'}.  KvidentI 
the  incri'asc  jiad  occurred  ahout  the  sanu'  time  as  the  passa'.^e  of  t'oo.' 
from  the  stimiarh   into  the  duodenum.     Thes(    I'acts   indicate  that   ure:i 

formation   in  the  liver  Ik mcs  slimulated  hinjr  Ixd'orc  the  other  tissue- 

sucdi  as  ti,.-  museliN.  have  iiad  lime  lo  lake  up  ilieir  full  <|uota  of  amiie- 


!    i 


Till;    MKTAHOI.ISM    OF    l'U()Ti:IV 


Gil 


ids.     Duriiiu'  •li'.'cstidii   ol'  protein   tl:o   livor  docs  iint   niipojir  to   Wiiit 

('(•(HIM'  sjitiiriitcil  witli  iiiuino  iicids  l)cfor(>  it 


Mitil  tilt'  otliiT  tissues  lune  li 

li  '_'iiis  to  destroy  tlie  nniii'ei'ssiiry  excess  liy  eoiiversion  into  uroii;  on 
tlie  eoiitr;iry.  tliis  process  sets  in  Avitii  the  very  first  installiiieiit  of  niiiiiio 
;i<-id  tlint  reiiclies  tlio  liver  liy  tlie  portiil  hlood.  Tliis  eoiieliision  is  in 
li.'iniioiiy  witli  tlie  well-estiiMislie.l  fiief  tlmt.  wIiimi  proi,.ii,  is  <riveii  to  a 
stjirviiiir  aniiiiiil.  tlie  irre.itei-  part  of  its  iiitro-;eii  is  soon  ey.erote.l  a's 
urea,  leavinu'  only  a  siiial!  fraction  to  lie  used  I'or  ichnildiiij;  tlie  wasted 
tissues   (  see   pa'je  (p-l.'i). 

The  amino  acids  that  are  alisorlied  Iiy  the  extrahepatic  tissues  Iiceomo 
very  (|'dekly  eoinerteil  into  t'oniiecl  inoteiii,  as  is  evident  t'roin  the  fact 
that  the  eoiiceiitration  of  free  amino  acids  in  the  tissues  of  an  animal 
diiriiiir  alis(irption  of  jirotcin  is  not   perceptiMy  u'fenter  than   in  those  of 

a    fastinu-  animal,  and   the  (piestion    remains  to   In nsidered.    Wlnil    hr- 

r„,ti(.i  of  till  priiliiii  llnut  fiiniiiil.'  The  answer  is.  that  it  is  iiradually 
used  up  in  the  iiietalioiic  processes,  so  as  to  iilierate  airaiu  the  amino 
:;cids,  which  add  themselves  to  those  alisorlied  from  the  intestine  and  lie- 
eome  used  atrain  or  csereted,  aecordin-r  to  the  demands  of  the  tissues  at 
the  time   for  amino  acid. 

Tlii^  1" '•■^  "f  liluu-ation  of  amino  acid   from  the  lireakdown  of  Iiody 

protein  "roes  on  of  course  irrcsjiectix  e  of  alisoriitioii  of  a,!iino  acid  from 
ilii'  intestine.  It  uoes  lui,  for  exampli-.  diiriii!.'  staivatioii;  indeed,  in 
this  eomlition  the  pei'ceiit.iL'-e  of  free  amino  acids  in  the  muscles  is.  if 
anything,  soiiieuliat  hidiei-  than  that  oliseiveil  in  an  ordinarily  fed  an- 
imal. Ill  starvation  also  the  iiiiH:ration  of  amino  acid  is  <zi>\\\>x  on  amoinr 
the  various  oriraiis.  of  whicdi  those  whose  activity  is  essential  to  the 
iiiainteiiance  of  life,  such  as  the  heart  and  the  ri'spiratory  miisides,  are 
supplied   with  amino  acids   from   tissues  that    are  less  \ital.  sudi  as  the 

skeletal  UiMseles  I  see  pairc  -"ifiS  .  Thc>c  cxpcrililclll  s  further  show  that 
I'ree  amino  acids  can  not  serve  to  any  si;;iiiticant  extent  as  feod  re-er\es 
■I  the  same  \\a\  as  -ilycociui  and  fat.  If  amino  acids  wire  of  value  as 
looil  reserves,  we  should  expi'ct  the  st.ire  of  Iheiii  to  lie  depleted 
iiy  starvation.  .\s  to  how  loim  a  pei-iod  of  time  elapses  hetwccui  the 
irieorpiM'alioii  of  the  alisorhed  amino  acids  into  tiss\ie  pidtein  ami  their 
>ulise(|uent   lilieiation  airain  liy  autolysis,  we  are  cuiliieU-   iirnorant. 

The  researches  which  we  h;i\e  .jiist  lii>en  coiKidiTin^-  do  not  throw  any 
'i-ht  oil  the  nliiliri  nihil  of  ililfiniil  jirnliiiis  in  liism  im  fnhull^ni. 
They  do  lint  inform  Us  as  to  which  of  the  amino  acids  must  lie  alisorlied 
!ead>-maile  from  the  divested  food,  and  which  of  thiuii  i  i;i>-  lie  <lispeiised 
with  since  the  <iruanism  can  iminnfacture  them  for  itsel-  We  !<iio\v  that 
the  hiirher  animals  can  s\nthesi/e  some  :iiiiiiio  aciils.  ^i|c!i  as  <r!\  eoeoll. 
hut   not   others,  such  as  tryptophane;   luit    which   amino  jn'ids  lulonur  to 


l!i 


.1 


!l    'i 


'ii 


612 


MITAltOI.lSM 


tlu'  kI.vcocoU  ami  «liicli  to  tlic  ti\  ptopliaiip  Kfoups,  can  not  as  \ii 
be  definitely  stated.  Tiie  invest ifjat ion  of  this  proiiJeni  has  to  be  undci 
taken  by  expeiiinents  ot"  an  entirely  <litTerent  type — namely,  by  observii- 
the  welfare  and  j;ro\\th  of  animals  fed  on  proteins  of  varying  amiii.. 
aeid  eomposition.  A  full  diseussioii  of  these  experiments  is  tjiveii  in 
the  chapters  on  Nutrition  and  (irowfh. 


}  H\, 


■■■  I 


ClIAl'TKR  LXIX 
THE  MKTABOLISM  OF  PROTPHN  (Cont'd) 

THE  END  PRODUCTS  OF  PROTEIN  METABOLISM 

Introductory.— So  far  we  liavo  approached  the  problem  of  protein 
metabolism  by  studying  the  behavior  of  the  absorbed  products  of  pro- 
tein breakdown,  and  we  have  seen  that  these  become  gradually  assimilated 
by  the  tissues  and  used  by  them  in  their  metabolic  processes.  We  have 
been  unable,  however,  to  ofTer  any  facts  re<;ardins  the  exact  chemical 
changes  which  each  amine  icid  undergoes  during  this  process  of  tissue 
metabolism.  At  first  sight  it  might  appear  an  easy  matter  to  collect 
such  information  by  direct  examination  of  the  tissues  themselves,  either 
by  searching  in  them  for  amino  derivatives  which  might  be  derived  from 
absorbed  amino  acids,  or  l)y  studying  the  changes  which  occur  when 
the  amino  acids  arc  sub.jected  to  the  action  of  the  isolated  tissue  en- 
zymes that  must  be  responsible  for  the  change.  Such  methods  of  in- 
vestigation are,  however,  fraught  with  technical  difficulties  so  great  that 
very  little  can  be  Icarnetl  from  them,  and  for  the  present  at  least  we 
must  be  content  to  j)ieco  our  information  together  fiom  facts  derived 
by  less  direct  methods.  Such  a  nietluMl  is  offered  by  investigating 
the  behavior  of  the  end  i>roducts  of  protein  metabolism. 

The  main  end  product  is  vrni  alor.g  with  traces  of  its  precursor  a»i- 
nwnia,  but  these  arc  not  the  only  ones,  for  some  amino  acids  after  being 
incorporated  with  the  tissue  proteins  lircak  down  into  products  that 
are  no  longer  members  of  the  amiiio-acid  scries,  although  they  may  be 
closely  related  to  certain  amino  ocids.  Such  substances  arc  mafiiir  and 
its  anhydrid  creatininr.  A  part  of  the  amino  acids  during  their  pres- 
ence in  a  free  state  in  the  blood  may  also  be  excreted  unchanged  by 
the  kidne,\ .  Our  list  so  far  therefore  includes  iirca,  ammonia,  creatine, 
creatinine,  and  annuo  nitrogen,  of  which  the  last  is  usually  included  in 
metabolism  investigations  in  the  fraction  designated  undetrrmined 
nitrogen. 

Another  group  of  closely  related  substances  coming,  not  from  the 
general  protein  metabolism  of  the  tissues,  but  from  the  metabolism 
which  is  peculiar  to  the  nuclei,  consists  of  the  so-called  purine  bodies. 
Furthermore,  so  as  to  serve  as  a  check  on  results  obtained  by  examining 
these  nitrogenous  metabolites,  it   is  impoi'tant  to  observe  the  manner  of 


w 


I" 


«llp.p 


T 


•111 


mi:tai!(ii.ism 


i;1  ill 


oxcii'tioii  ,,|'  tlir  siilpliiir  ictv   i<(  tin-  |initciii   hioIitiiIc.   I'or  ii    \\\\]   1., 

r.'iriciiilMTi'.!  tli.il  it  is  ill  |. ml. -ill  nldii.-  ili.it  siiliilmr  is  iisii;ill.v  l;ilv,.|i  iii 
till'  .'iiiiiiiiil  Imi.Iv.  Till'  cMTi'tioii  .if  snlpliiir  tliiiclorc  riiiis  iinirc  ,ir  !.■- 
|>iil;illi'i   witli    tlic   iiil.'iisity   i.t'   inotrin    iiu'liili.ijisii). 

Al'l.T  s.'Iri'tiii-r  till'  I'liij  |.ri:.Iiii-ts  tlijit  iiri'  most   lik.'ly  I..  I r  si-ni- 

ic-.'IIK-l',    till'    (ir-t    .|lll'stil)| Ilci'llls    ih..    ill it    ,,)■    ,.,.|,.1|    ,,f    tlii'lll    I'MMll.    : 

liiii-iiii;  Iwi'iit.s   roiir  111, Ill's  on  iliris  tlijit   .'III'  I'itlii'i'   rii-li   or  inpi.r  in   pi. 

ti'lll.       'i'lli'    iMPSsiliJIily    ,,f    i-iiii.liirtjiiir    slli'll     ill\i'sti'_';itii(lis    nliviollsK     c|. 
lii'iKJs  nil    till'   Use   of  .|iiii'k   .'iiiij   yet    ii'linlili'   iiii'tliiiijs    fur   tin'   rst  iiii/it '..i. 
(>{  till-  iiitr.p.,'i'iiiiiis  iiii'tjiliiijili's.      Slli'll    iiii'IIkmIs   |i;ivc  lic'n    riiiiiisli,.,!    I.' 
till'  liiiilisf.'ikiii!,'  .'iiiil  riiicfiil   ui,il<   ,,r  Kiiliii.  jiii  ,.xiiiii|il,.  of  wliiisc  I'l'siiil. 
iii'i'  iri\rii  ill  till'  ;ii'r(iiii|);iii.\  iiiji  tnlili'. 


X'lihiini'  I'l'  iiriiir 
'l'nt;il   nitrogen 
l'ri':i   iiitiii;.'!'!! 
Aniriiiiiii:!    iiilii>:,'i'n 

I'lil-iwiil     Ilitli.;;iMI 

*'ri':tl iiiiiir  Mtltomti 
I'lnli'tt'Mriiin'.l    nit  iiwci 
Ti.tiij   SO 

Ill.M-Ullllir    SI) 

i;ilii'ii:ii  so. 

Ni'iilnil   SO,' 


xitkik,i;n  iMiii  iiii  r 

Ii'i.s    i;i;nii'4 

11.7     miiiii-  s;.,-,-; 

II. til  L'liiMi  :;.ii', 

II.  IS     ;;|;ini  |     |  '  ; 

i'..''S  L'liim  ;;.!•'; 

I'."".')  L't.'ijM    -  l.'.i' ; 
.■;.ill    -rams 

.'M'7   ;.:r:iins  '.ni.H' ; 

11.1:1   t;i!ilil     --  .'i.L''; 

ll.1><    UHII"      r  r  t.S'; 


NlTlltXiKN   I'liOK     I'll 


;i.ltll  ;ii:(i(is 

L'.-'ll  uraiiis  -r  UI.7' 

II.  IJ  ii'Miii    n    1 1.:;- 

ii.iii"  '.:iaiii   --   :.',."p' 

li.llii  i;iiiiii    -.-  17.1' 

II. J7  t; I       -     7..:' 

ii.7l>  -1:1111 

II.  Hi  ;;ii(ni     __  lin..".' 

I'.K'  ;;iiiiii    --  i:;.L' 

ii.:;ii  uKini    —  L'li.:; 


rKiiiiii.i 

TIlc   fii'iicfiil   I'liiicliisii.iis   \vliicli    iMJiy   In-   .Irjiwii    t'i'iiiii   tlii'si'   icsiilts   ;ii. 

!1S    t'oIIllAVS: 

1.  AN  itii  ji  iirotiin  I'ii-li  .lirt  imii'li  nioii'  iiiiiu  is  cxcri'tcil  in  twi'iit^ 
four  liiiiii's  tliiin  witli  niu-  tluit  is  |.i'nti'iii-|HMii'.  K-,  iiL-iitly  tlic  iiiti'.i-i'iiou 
ini'tiiliiiliti's  iirf  iis  (liiiii'tii's. 

•_'.    Till'    tut.'ll    111'    jiiisolnti-    ,'iiiiiiiiiits    of    llitloy;!'!,    iiiiil    of    Jill    tlic    i.tlh 

iiiti'ou'i'iioiis  iiii'tjilii.liti's.  siivi'  I'li'jitiniiir,  1 mi'  iliiiiiiiislii'il  .liii-iiiu  tl;. 

st;ir\;itinii   iMiioiI.     The  s.iiiif  is  tnii'  of  tln'  siilpliiir  il.M'iv.itivt's.  cx.ii. 
ill  till'  I'.'is."  of  till'  iii'utnil  siiliiliiir,  wliii'h  lu'lmvi's  liUi'  cicjitiiiiiu'. 

:'..     Till'  lll'I'lCMSI'    ill   till'   |)oltili|l   111'  llilr'oircll   CMTctl'.l   ;ls   llll'll    Is   I'l'ljl  t  i\  ('1 

Kii'.it.T  lli.'iii  tlic  ilccic.'isc  ill  totiii  iiitriiri'ii.  this  liid  l.cinir  sliown  in  tli- 
tiililc  liy  till-  pci'ci'iil.'itrc  lltiin'cs.  which  were  sccincij  liy  I'lili'iil.-it  in 
till'  iii'opiiitiiiii  of  nitrn!.'cn  in  the  vjiiimis  siilistinccs  ;is  a  peieent.iL: 
of  the  total  nitroLren  excreted  diirin;,'  the  iicriods.  The  inoru'aiiic  sir 
Iiliatc  heliavi's  in  a  iiianiicr  similar  to  the  urea  that  is.  the  pereciita'j 
of  total  snlphale  excreted  in  the  inoru'anic  I'oriii  Iiccomu-s  iiiucli  Ic- 
diirimr  starvation. 

4.  The  relative  anioniit  of  all  the  otlier  nitroirciions  iiietaliolitcs,  :i 
well  as  that  of  the  clheical  and  neiilral  siii|ilia1es,  iiccoiiics  inercasi  • 
(lufinf;  sfai'Viition. 


i\i 


^1:  :i^ 


Tin:    MIT.MIOI.HM    OK    I'HiPTIIV 


(;i:. 


Tlir  llici^l  slliliilii.'  ii'slllls  u(  tlic  iihiivi'  in\  I'st  i'.'iltiiiii  ;iic  tliMf  crc-iliiiinc 
IcMl.iilis  Miirliii|ii;iil  illllilii:  vt;ii\;iticill,  lull  tliiit  llli'll  Iiimmpiucs  rclillivcly 
iin-rcji-^i'il.  'I'lir  Iniiiii'i-  iiiiisl  III'  iji'iivcil  I'nnii  iiii'(;iliiilii'  iiroi-i's'.i's  truiim' 
nil  in  llif  IInsih's  iiiijcpi'iiilciilly  ol'  llic  Mip|il,\  nl'  rniiilstiilT  i-jiiTicd  to 
llii'in.  w  lii'ii'iis  tlic  hiltiT  liHisI  (Icpi'iiil,  if  nut   I'litiii'ly,  vcf   very  l.'irjrt'ly. 

■  Ill  till-  |i|-oli'iii  fiiiiti'iil   of  till'  f I.     Cii'Mliiiiiii'  iii;i>    tliiTi'fiii-i'  III'  I'lillcil 

Mil  cllil  prmlllrt  of  I  iiihiiii  nulls  liictjiliuli-lil.  .'linl  llli'M  nil  cihI  pliiillli-t  of 
I  riiiii  nulls   liii'l.'ilioliMii. 

()tlii'r  iiii'tiiliiiliti's  njiiiii'ly,  .■miiiiniii.-i,  iirii-  in-iij  inul  tlii'  iiiuli'tcriiiiiii'il 
iiitroircn,  iis  wrll  jis  tlu'  ctlicri'nl  Milpli;ili's  must  ii'pri'M'iit  iifoccssi's 
of  iiii'tnliiilisiii    tlijit    .-HI'   |i;iitly   cxo'j'i'noiis   ;iiii|    ii;iilly   I'liiloirciioiis. 

Il;i\iir_'  iii.iili'  oiiisi'hi's  .•ii'i|iijiiiiti'il  uitli  tlu'  iii'inTiil  n.itiiic  of  tlu' 
rlijiiitri's  tli.-il  oi'ciir  ill  till'  niti-iit;iiiiiiis  iiict.-ilioliti's  wliin  prolciii  mi'lali- 
iilisiii  is  stiiiiiihitcil  liy  tlic  tiiUiiiij:  of  fooil  or  ili'in-rsM'il  liy  st;nv;itiiiii. 
\M'  iii.'iy  now  pioci'iij  to  t;il<i'  up  tmi-Ii  of  the  iiu'tiiliolitrs  si'pii iiiti'ly  jiml 
SIT  \\liiit  otliiT  infoiiiuilion  r.iii  lie  olitiiiiii'il  ii'iriiiiliiiy;  tlirir  siiiirci-  iiiul 
oriirin   in  tlir  iiniiiial  lioily. 

UREA  AND  AMMONIA 

{•"or  \ai-ious  i-casoiis  it  is  iiiipiulaiit  to  coiisiili'i-  llusi'  1 -a  o  iiii'taliolitcs 
liiiri'tlicr.  Diiiiiiji  tlu'  inti-iini'iliary  iiictaliolisiu  of  tlic  majority  of  tlic 
aiiiiiio  aciils,  tlic  amino  u'loiip  liccomcs  Inokin  otV  as  ammonia,  A\liicli 
immediately  coMiliiiics  w  itli  tltc  a\ailalilc  aciils  to  foini  iiculial  aiiimonimii 

salts.     Tlic  most  availalilc  aciil   for  tliis  |)Ui-posc  is  carl ir  aciil :  tlicrc- 

I'oi-c  ammonium  carhonatc  is  formcil  in  laryc  amounts.  A  small  proi)or- 
tion  of  tlic  aiiiMioiiia  ma.\  coinliinc  witli  otlicr  i'ciil  railiclcs.  sudi  as 
rliloriiic.  to  form  ammonium  diloriilc.  Tlic  fate  of  tlicsc  two  t.\  pes  of 
salt  is  very  ililVerciit.  Tlic  aiiimoniiim  carlionatc  liccomcs  ipiickly  trans- 
formed into  urea,  wlicrcas  tlie  amiiiiuiiuni  cliloride  is  excreted  in  tlie 
urine.      Tlic   process  of  iirca    formal  imi    may   tlicrcfore    lie   considered    as 

liavinu'  the   function  of  pre\ciitinir  tlie  ,- niiiulaliou   of  ammonium  cai-- 

lionatc  in  tlic  animal  lioi|.\-.  It  is  the  means  liy  wliicli  :\  liarnifnl  sulistaiicc 
is  con\erteil  into  an  iiiiineiiiius  siilislanec  a  diloxicatioii  process,  in 
iitlicr  words. 

IJcLrardiiiir  //"  iiiiliin  nf  lln  rln  niiinl  jinn-i  ss  involved  in  tliis  trans- 
fiirmatioii  of  animnniuni  carliunate  into  urea,  rcfcrciiec  to  tlic  fornnilas 
lielow  will  sliow  lliat  the  aiiimoiiiiim  carlionatc  that  is  formed  liy  the 
union  of  c;irlioiiie  acid  with  aiiimoiiia.  liy  losinir  one  iiioleenle  of  water 
iiecomes  aiiimnniniii  cariianuiie,  which  liy  rcjielitioii  of  tlie  iiroecss  lie- 
comos  transformed  into  urea: 


'is     I 


il 


f 


(if 


i:    ■! 


i    i 


!  11'  ; ! 


616 


MKTABOr.ISM 


nil 


OMI, 


r«)  L'NII,r=fr() 

\  \ 

oil  OMi. 

(l'!llli"Mi.'  ClIlirilM  ('•ilDIIliilliuill 

iii'i'l)  iiiiil  i-MtlH.iiiilri 


II  <);=?»•() 


/ 


nxH. 


MI, 


If.O  — (Y) 


Ml, 

f.'iiiinioiiiiiin 
I'firliMiiijiti'l 


\ 


Ml 


f  iiii':i ) 


Soino  of  tilt'  iiira  iii.-iy  conic  fmin  iiictjiliolic  j)roccss(>s  of  iiii  nntircl', 
•  lilTcrciit  fvi)c.  f)iio  of  ilics.'  at  least  is  known;  iianicly.  tlie  splittiiiirntT 
of  mm  from  ai  (.niiinc.  wliidi  it  will  !).>  rcinoinlicrcil  is  j.'iiaiii(!inc-aniiii.i 
valerianic-  acid  i  s.'c  lufrc  tllC).  An  cn/ymc  calld  arjrina.sc,  liavinj;  tlii. 
action,  lias  liccn  ivolatcil  from  varions  orirans  and  tissues.  The  (liamitio 
valerianic  acil,  or  ornitliiiic.  wliicli  remains  after  tlie  urea  is  s])lit  otT 
may  I)e  further  used  In  jijotein  melaliolism.  Tlio  reaction  is  .shown  in 
the  followini;  ci|iiation: 

Ml,     TMI     (11^     (11.     Cir^  CIINII      conll.lio 

^'^f  (■.■ii^'iniiii''! 

Nil      CO 


MI 

luriM  I 


Nil.    <ll     <ll.    ni.    riiMI,    COOK 

fninilllilic') 


On  an  ordinary  diet,  as  we  have  seen,  a  man  cxi-retes  somewhat  more 
than  ftO  |)er  cent  of  liis  total  nitro^'cn  as  urea  and  aiiont  :{  i>er  cent  .is 
ammonia,  the  remainder  of  the  nitro<.'cn  apjiearin-;  in  the  other  iiitro..' 
enous  metaliolitcs. 

Influence  of  Acidosis  on  Ammonia-Urea  Ratio.  Tt  sometimes  liap|)ciis 
that  a  larire  proportion  of  the  ammonia  is  not  convert.'d  into  urea,  Imii 
is  use<l  for  the  ])uri)ose  of  iieutralizin<r  aliiun'mal  acids  present  in  tlir 
oru-anism.  AVhcn  mineral  acids  arc  fiiven  to  an  animal,  or  Avhen  acid- 
are  produced  in  the  orjranism  itself  liy  some  faulty  type  of  metaholisiii, 
tlie  ammonia  exi-iction  hy  the  urine  immediately  rises,  In  diabetes,  for 
example,  where  consideralile  (piantities  of  /i-o\yl.ntyrie  acid  are  l)ro 
dueed  (se..  pajfe  (iS.!).  j,  dccM.ied  inerci.se  in  til.-  amiiioiiia  ex.-retion  I,; 
tlie  urine  is  ohs.-rved.  A  mihler  type  of  acidosis  may  also  he  induced 
in  normal  persons  Iiy  wi;]iholdin>_'  carliidiydrates  from  the  diet,  and 
here  ajrain  the  ammonia  exc-retion  is  relatively  increased. 

In  such  eases  it  is  ipiite  evident  that  ammonia  is  used  as  <ni  nlkaliii. 
risrrvr  of  the  Ixidy;  that  is.  as  ;i  sulistance  which  is  eajiahle  of  prevent 
iuK  acidosis  liy  nentralizin-r  the  aeids.  It  does  not  appear,  however, 
that  all  types  of  acidosis  entail  the  utilization  of  ammonia  as  reserv.- 
alkali,  and  an  inriea.M-  in  the  relative  amount  of  ammonia  in  the  uriin 
does  not  necessarily  indicate  a  condition  of  acidosi.s.     In  the  pernieious 


THE    MKTABOMSM    OF   I'HOTKI.V 


617 


vomiting  of  prcKrinncy,  f(ir  example,  n  relatively  high  excretion  of  am- 
monia has  heon  IoiiikI  associated  witli  no  jjreater  a  decree  of  acidosis,  as 
determined  by  the  power  of  the  i)lasma  to  ahsorl)  carbonic  acid,  than  in 
normal  eases  of  jjreffiianey. 

Influence  of  Liver  on  Ammonia-Urea  Ratio,— Experimental  Obierva- 
tiona:  (1)  Rkmovai-  ok  Livkk.— There  are  several  facts  which  indicate  that 
iilher  c'luscs  than  acid  j  idiiction  may  interfere  with  the  conversion  of  am- 
monia into  urea.  AVliat  arc  these  causes?  Since,  as  we  have  seen, 
the  liver  is  the  ors;an  which  most  actively  converts  amino  acids 
into  urea,  it  would  l)e  natural  to  expect  that,  when  the  functions  of 
this  opfriiM  were  interfered  with,  relatively  more  of  the  nitrosrcn  excre- 
tion would  occur  as  ammonia  and  relatively  less  as  urea.  In  order  to 
determine  the  exact  sifrnifieance  of  the  liver  as  a  urea-forminjr  or-fan. 
two  types  of  investiiration  have  been  used:  namely,  (1)  observatiim  of 
the  channes  produced  in  the  animonia-urca  ratio  in  the  urine  by  partial 
or  total  removal  of  the  liver;  and  (2)  observation  of  the  urea-fcirmiri!.' 
power  of  a  liver  perfused  outside  the  body. 

To  remove  the  liver  from  the  ciiculation  the  portal  vein  is  brought 
in  apposition  with  the  vena  cava,  tlic  two  are  sewed  to<rether,  and  a 
passatre  opened  between  them,  after  wliieli  the  portal  vein  is  li<rated  above 
the  anastomosis  (forminrr  the  so-called  E<'k  fistula V  The  portal  blood 
then  passes  directly  into  the  vena  cava,  and  the  liver  is  now  supplied 
only  by  the  hei)atic  artery.  The  animals  live  for  a  considerable  time 
after  the  operation,  and  the  urine  fre<piently  contains  relatively  less 
urea  and  more  ammonia  than  normal.  The  results  are.  however,  not 
nearly  .so  stril<inp  as  would  be  expected  if  the  liver  -were  the  main  seat 
of  urea  formation.  The  experiments  have  nevertheless  brouirht  to  light 
a  fact  of  considerable  clinical  interest- namely,  althou^'h  the  animals 
may  thrive  if  kept  on  a  diet  not  ( ontaininsr  an  excess  of  flesh,  they  im- 
mediately bci;in  to  develop  peculiar  symiitimis,  not  unlike  those  of  ec- 
lampsia or  uremia,  when  they  are  fed  with  lar^rc  airiounts  of  flesh  food. 
Most  of  the  synij)toms  can  be  referred  to  abnormal  stimulation  of  the 
I'cntral  nervous  system,  and  examination  of  the  urine  has  shown  a  lari;e 
increase  in  the  excretion  of  ammnnia  and  a  ehange  from  the  normal 
acid  reaction  to  an  alkaline  one. 

At  one  time  it  was  assumed  that  the  to.xic  symptoms  Avere  caused  by 
the  presence  in  the  blood  of  ammonium  carbamate,  since  large  quantities 
<if  the  calcium  salt  of  this  substance  could  be  separated  from  the  urine. 
It  is  now  known,  however,  that  the  ammonium  carbamate  is  present  only 
because  of  the  excess  of  ammonium  carbonate,  the  two  salts  existing  to- 
irether  in  solution  acccwdinir  to  the  laws  of  mass  action.  That  the  intox- 
ication is  not  due  to  ammonitnn  carbamate  does  not  exclude  the  pos- 


i     I 


!       ■  it 


i 


1^  III  ■! 


'i'   ' 


(IIH 


mi:t.\imii.ism 


siliilily   tli;il    it    liiiiN    lit'   i|u(.   I,,   iitllllinliiii    itscll',   iiltlliMluh    i)    is   lili.r,.    lil,, 
lll.lt    olIl.T    nil  ll.«;.'ll. Ills    IMClill.olit.'S,    |irili|||,M'l|     wllrll    l-MM'SH    <l|'    llcsh     I ' 

is  tiikfii,  nil'  till'   i')'s|iiiiisili|i>  iiifi'iits. 

11"  till'  liviT  is  iiiliiihi  niiKii;,!  \,y   lii;iitiiiy  tlif  lirpalii-  iirti'i-i.-s  in  ,i - 

.llllliiill    witll    till     Ki-k    listlllii.    il     liiori'    pi Illii-t'il    ilrrri'flsi-    in    lllfji    ;ih  : 

ni.TciM-    in   ;iniiiiiiriii iMir  iluiin'r   tin-   slimt    iicriml    i.f   tiiin'   tliiit    11 

iiniiiiiil  siirviM's  tln'  ii|M'i'iitiiin. 

Till'  iisniN  ulisiTM,!  artif  ilir  ii'innv.-il  i.r  iliininiitinn  ol"  liver  t'nmMi'.r. 

''"'   ' "I'  "'I'll   "Ili'T  \isc'iTii   ail'   niiiKVi'il    iV.ini   tlic   iininnil,   ulihl. 

W'.nlil  ill  least  t.'ii.l  to  itiilieale  that  the  liver  is  very  inipi.rtaiit  in  Up 
liiaiiiifaetnre  of  tiiea  oiil  of  aiiinionia.     Tliis  does  not.  however,  warrant 

the  eonrliisioii  that  the  liver  i:.  the  only  pli ill  Hie  animal  l.oily  in  uhi.-i, 

sllell   a    pi'oeess  oeenis, 

III  eorroIioratioM  of  these  oli- ervat ions  on  iiiaiiiinals.  it   may  he  of  n. 
terest  to  note  that  iih,  ii  II,,   I,,;  r  Is  r,  „i„i;  ,1  fniin  h,r,ls.  whieli  is  a  enm 

p.'iratively  simple  operation  on  ; iint  of  a  natural  anaslomosis  helween 

the  portal  ami  renal  veins,  there  is  a  markeil  ileerease  in  the  exeretio- 
«f  iirie  aei-l  ami  a  eorrespomlin..'  ineiease  in  the  exeretion  of  ammoni, 
•  liliiiiir  the  twehe  hours  or  so  that  the  l.irils  survive.  In  hinls  an! 
reptiles    urea     is    exerelei!    as    lirie    aei.l,    liein..'    priHlnei'iI    liy    a    s.Mitllcli' 

'" ^^    ''I    "l''   li^il'    's<>e    pa..',.   (I-M).      The   ehamjes    ill    this   experiliienl    ,11. 

"''  ' siileral.le    maj-nitiiile;    thus,    liefoie    the   operation    the    aiiioinit    .  ' 

aiimioiiia  iiitro._'en  relative  to  total  Milio.,'eii  has  lieen   fomi,]  to  vary  h. 
tweeii    II)  ami    IS   per  cent:  after  the  operation    it    may   he    iiiereaseil    • 

hetweeii  4.")  ami  llll  per  cent.     The  iirie-aeiil  nitroue riiiall.x    varies  \.. 

twoeii  lil)  ami  711  per  cent  of  the  total  iiitio._'en:  after  the  operation  it  im,i. 
fall  to  between  :!  ami  Ii  per  eeiit. 

Ill  aniiiials  with  an    Kel<   (isliila   ami   with   the   hepatie  artery   lifjatr.! 
an  iiierease  in  the  urea  output  oeeiir>!  when  amino  aeiils  are  in.jeeteil  iimh  ' 

the  skill.     This  result  eoriohorates  tl iieliision   that   the  liver  eaii   ic 

alone  he  resi)onsil)le  for  the  eonveisioii  of  ammonia  into  urea. 

r2)  I'lHKi  SKIN  OK  ()E{(iA\s.  This  iiietlioil  eoiisists  il;  reinovin.,'  the  o; 
iraii  into  a  warm  ehamher  or  hath  ami  perfiisiiiir  it,  t|iioiii.'li  eaiiniil 
inserted  in  its  main  arl.'ry  and  vein,  with  a  solution  of  delihrinated  l.h". 
or  of  delihrinated  lilood  mixed  with  s.iline  solu'ion.  The  perfiisi.. 
liijuid  is  kept  at  liody  temperature  and  is  saturated  with  oxyireii.  I', 
means  of  a  pump  it   is  made  to  eireiilate  in  a  pulsatile  How.  and  the  to!; 

amount  of  urea  or  other  iiie,ahoIite  in  t| ireiilatin.,'  Iliiid  is  determiii. 

Iiefore  and  after  the  lluid  has  I.eeii  eireiiiated  several  times  tlirousih  tli 
(ir-raii.     Wh.'ii  the  liver  is  iierfiised,  urea  ^'radually  aeeiimulates  in  tl 
(hiid.   parlii-iilariy   afi.r   the  addition   of  one  id'   its    known    jireeursors 
fur  example,  ammonium  earhonate.     When  other  orjiaiis  or  viseera   ai 


tin:  mi  •iaim>m>\i  ay  iki.ii  is 


(i1'> 


pcifiiscd,  nil  iiiTii  is  rmiiH'il.  'I'lic  cviilrini'  sliows  ilmt  tin-  liver  is  nil 
illl|)iil'tilllt    M'llt    III'   liri','|    I'll!  Mini  lull,    lull    lint    lirri'ssjii'ilv    tlllll   mIIii'I'   nruillis 

lilt'  iiiiiiiilc  III  ''iiiiii  it  ill  till'  iiitiirt  aniiMiil.  I'ur  tlini'  iiic  iiiaiiy  soiim^ 
iif  iiiiirriiriifii  in  /n  rfiisimi  i  i  /n  rinn  nl^.  Kmm  lliiiiiifli  we  cxcri'isc  tlii' 
'.'i-i'iifi'Mf  fiiri',  \\i'  ciiii  lint  liiipi'  til  iiiMiiilMiii  till'  iiiirjiii  ill  .illicr  fliiiti  a 
slowly  ilyiliit  I'liiHlitiiiii,  It  is  riTtiiiiilv  far  li'iimv  nl  rinin  tlic  liniinal 
state,  as  is  ii'vcali'il  imt  milv  liy  liislnliiific  fxaiiiiii.iliiiii,  liiit  liy  llii'  fact 
that  I'lji'iiia  aliimst  iii\aiial>iy  sets  in  ami  tlic  liliiml  m'sscIs  liccniiir  rx- 
tri'iiii'ly  ('iiiistrird'il.  thus  lui'i'ssitalinir  a  irfailiial  iiiri'i'asi'  in  llir  pcr- 
riisinii  pii'ssiirc  as  the  pn  riision  (foes  tm.  Fiiitliriiiinii',  the  inu'aii  liriii',' 
isiilalcil  t'l'olii  flic  iH'rvoiis  s\sti'i|i,  llicic  eaii  lie  iin  niiilriil  of  tile  rela- 
tive liliMnI  supply  of  ilitTeieiit  parts.  In  the  iiitaet  animal  the  eireiila- 
tioii  is  iiioii'  or  less  ilistrilmteij  aeeoiilintr  to  the  partieiilar  needs  of  tin' 
ilill'i'ient  xisiMl'a.  aiiii  slleli  eniiilitionx  oliviolls'y  eail  not  lie  sillllllateil  in 
a    perfusion    esperiiiieiit.      Another   olijeetioii   ilepeinls   on    the    fact    ihaf 

the  uell  liein'_'  of  tl ruiins  in  the  intaet  animal  is  larjrely  ilepenilent  on 

hormones  eoii\e\ei|  to  tlieiii  fioiii  otln'r  organs.  Siieli  hoiniones  are 
fieipienll\  ipiite  lahile  in  nature,  ami  soon  ilisappe.ir  fioni  the  perfusion 
llui.l. 

Notw  ithstamliiii;  these  nlijeetioiis.  there  ean  he  no  iloiilit  that  many 
of  the  fiinetioiis  of  ail  oru'an  are  retaineil  iiiiieh  Imi'.'er  than  they  woiihl 
he  if  the  oriraii  were  not  peifiiseil  ;  foi'  example,  the  eonl  raetility  of  the 
iiiilsele  or  the  ))ii\\er  of  foriiiiii','  urea  in  the  li\er.  I'eri'usioii  exiierimenls 
.•ire  of  \alue  therefore  when  they  yield  positive  results.  Neiralive  re- 
sults may  indieafe  either  that  the  oiyan  does  not  perform  the  particular 
fiiiietiiin  that  is  lieiiii;  iii\  I'stiu.ited  or  that  it  has  lost  this  function  as  a 
result  of  partial  death.  That  a  perfiiscij  muselc  retains  its  power  of 
rontractioii  does  not  iieecssaril.v  indicate  that  it  maintains  all  of  its 
iiietaholic  functions:  neither  does  the  fact  that  the  liver  forms  urea 
prove  that  it  is  eapahle  of  pcrt'oniiiii','  its  other  functions.  It  is  eas.v  In 
show  Ihat  the  li\er  dies  pieeenical;  some  flllietioMs,  such  as  };lyco(;ell- 
formafion,  die  eari.\'.  while  others,  sinh  as  urea-formation,  ri main  for  a 
loinr  time  intact.  77//  I'si  of  pi  rfiinlnii  i  ipirhm  nta  for  tin  stltliiiij  of 
'liiisfions  of  nil  tiihollsni  slioiilil  I  Ik  ri  Ion  uhiniis  hi  n  ni  run  fiillif  coii- 
Irolliil  mill  III  VI  r  iisnl  ns  lln  siili  Inn  of  niili  iin  on  iihicli  to  hiixr  iiiii>or- 
liinl  conrliisions. 

{'■])  llefore  leaving  this  sulijeet  it  lita.v  lie  udl  to  |iiiilit  ollt  that  the 
inelhod  which  at  first  siirhl  ini'rht  ajipear  to  lie  the  simplest  for  settlinj.'  such 
i|Ucstioiis  naincly,  thr  (in  mi  nut  ion  of  tin  inlUnriiui  nnil  oiitltoivimi  blood 
of  ililhrint  parts  or  oninns  -\s  not  applicilile  in  most  cjises  This  is  lie- 
eause  of  the  extremely  small  ehan^'es  in  concentration  which  may  occur 
even    !ilthoui;li   larjjre   amounts   of   the    partieul.ar   .substance    in   (luestioii 


M 


\U;\ 


ll 


II 


*!  ■ 


iil 


is! 


k 


\i\ 


w^ 


i    1 


W    ill 


r.2() 


MKTAHOt.ISM 


are  l)oiiiir  nliHoilml  or  prixlucp.!.  As  \y(^  nIihII  ncc  later,  thin  oriticiNni  .. 
particularly  appliciihlo  in  tlio  .-nv.  nf  sutrar.     Kvpm  .luring  the  injet.fi,,, 

of  considonililo  .|uanliti.>s  of  xuKar  int..  the  portal  vein,  no  diffor... 

in  pen-ontiiKf  ••an  hv  denionstrntcd  ln'tueon  the  Mood  of  the  two  mI.I.  ^ 
of  the  liver,  although  we  know  that  suKar  is  liein)^  retained  to  form 
ifl\ eopen.  For  the  same  reasons,  dilTer.nees  in  the  pereentaKc  amount  - 
of  amino  ai-ids  or  i.f  men  are  often  <iinieiilt  to  demonstrate  in  the  hlo-.l 
enterinif  and  leavinir  the  liv.r  even  when  we  know  that  lnri?e  qunntitir. 
of  them  are  lieini;  added  to  or  remove.!  from  it. 

Clinical.     Sinee  the  liver  is  an  iinportant  sent  of  urea  formation,  tl,,. 
•  luestion  arises  as  to  wheth.-r  the  relative  pereentajfe  of  urea  and  am 
monia  in  the  urine  will  heeome  altered  by  ilhrasr  of  the  llvrr.     Mal,^ 
ohservations  with  this  point  in  view  have  I.een  undertaken,  hut  it  cnu 
not  he  said  that  the  residts  are  very  strikinir.     In  extreme  destruefi.,i,, 
sneh  as  that  jirodui'ed  hy  phosphorus  poisoniriR,  there  may  indeed  !"• 
n  prent  inerense  in  the  relative  amount  <.f  ammonia  nn<l  n' decrease  in 
that  of  urea.    The  same  is  true  in  aeute  yellow  atrophy  of  the  liver,  in 
whieh  diseas..  the  nifrou'en  exerete.i  as  amm<min  may  amount  to  as  mini, 
as  70  per  eent  ..f  that  exer.-ted  as  urea.     In  milder  forms  of  liver  di> 
turl.anee,  however,  sueh  as  eirrhosis,  th.'  (inures  are  mueh  less  strikiir/ 
When   an    increased    anunonia   excrctim    is   observed    in    such    cases.    v\. 
must  he  cautious  in  drawinsr  the  conelus...n  that  it  is  due  primarily  |.. 
aholitioii  of  the  hepatic  function.     If  may  Just  as  well  lie  caused  l.y  tli. 
devel.ipnu-nt   of  acids   in   the  oriranism   that   require   the  ammonia   fm- 
their    neutralization.      It    is    sifrnifli-ant,   for   example,   that  consideraM- 
(pmntities  of  acids  are  produced  in  phosphorus  poisoninp. 

Althonjrh  the  urea  an.!  ammonia  excretions  heeome  altered  hy  extoi. 
sive  destruefior    ,f  liv..r  tissue,  it  is  a  remarkable  fact  that  very"  little  ii 
any  change  occurs  l„  fh,   nmim,  uilrwn  n.  either  of  the  urine  or  ..f  IIk 
blood.     In  experimental  necrosis  of  the  liver  prmluced  bv  chloroform 
or  by  phosi.horis.   it    is  only   in   the  latest   stapes  of  the   condition   an.i 
when  It  is  of  the  very  severest  type  that  an  nnino-acid  increase  has  be,  - 
found  to  occur  in  the  blood  and  urine.     The  conditions  seem  to  be  som. 
what  dilTerent  in  man,  abnormally  hi-:li  amounts  of  amino  nitrogpn  h.i\ 
iuK  bc.'n  observed  in  the  blood  in  a  considerable    proportion  of  patieii- 
with  impaired  liver  function.     In  very  severe  cases  of  diabetes,  for  .  v 
ample,  fig-ures  that  are  distin.tly  lii-her  than  norn),nl  have  been  observr! 
(Van  S'yke,  etc.K     In  e.-lamjisia  the  marked  i>atholo!ri,.  ehnnRes  in  tii. 
liver  niiiiht  be  exriected  to  be  associated  with  an  up.set  in  the  mctab. 
lism  of  amino  acids.     Losee  and  Van  Slykc^"'  have,  however,  recent  1 
.shoun  by  llic  most  accurate  methods  that  neither  in  the  blood  nor  in  tl 
urine  is  any  excess  of  amino  acids  to  be  found  in  this  condition,  alth.n--; 


Jul: 


Tin;    MKTMMII.IhM    (If    IKIITIIV 


»;l>i 


in  vasvH  1(1'  pfinicious  vomiiiiiK  of  |tnKiiiiiir\ ,  iIh'Ii-  «ns  a  relnlive  iii- 
iTPO!'"  in  tlu"  iiiiinioriiu  <'XciTtii)ii.  Wf  liiivf  iilrcntly  nccn  that  tliia 
iiicu-asi'  (li<l  not  hoar  any  n-lationship  to  the  ai-idahsorhinn  power  of 
the   hlo(Hl   plaxiiia    (see   pa-"  •117). 

The  importiituf  of  tin  kliliirim  in  nmorimj  llu  una  from  the  blood 
is  rt-ailily  seen  from  th<>  chaiint'  in  the  pi-rpcntasje  of  urra  in  this  fluid 
fiftiT  th(>  parfinl  or  coniiilctf  nninvnl  of  tho  kidneys.  Animals  sur- 
vivf  nephrectomy  for  about  three  days,  and  iliirinn  tliis  time  urea  rapidly 
aeeiimnlntes  in  the  blood  and  begins  to  make  its  apiiearnnec  in  the 
siiliva  and  the  intestinal  seerelions.  [n  man  also  where  the  kidneys 
are  extensively  diseased,  a  similar  aeenmulation  of  urea  oeeurs  in  the 
Idood,  some  of  the  excess  beiuK  Kot  rid  of  through  the  sweat  and  to  a 
certain  extent  through  the  intestine.  The  importance  of  eneouraging 
perspiration  and  a  free  movenn-nt  of  the  bowels  in  eases  of  nephritis  is 
thus  indicated.  It  must  not  be  eoneluded  that  the  aceumulution  of 
uren  'n  the  organism  is  the  direct  cause  of  the  symptoms.  I'rea  itself 
is  eomparntively  inert,  and  it  is  irenerally  believed  that  other  metabolic 
Iiroluets  with  which  the  urea  runs  parallel  in  amoiuit  are  the  toxic 
nu'ents.  Hewlett  has  found,  however,  that  very  l;irue  in.jections  of  urea 
•hi  produee  .symptoms  in  animals.'* 


^ 

t 

1 

1 

i 

T 


•i 


CIIAI'TKIf   I. XX 
TIIK  MKTAI'.OIJSM  OF  I'liOTKIX  (CmitM) 

CREATINE  AND  CREATININE 

Crfiiliiio  ami  .•rciilininc  iin-  very  ljirj.'.'Iy  |iniilii.'ts  of  (Middfrf'iK.iis  iin't.-ib 
nlisiii:  llicy  air  mainly  dciiv.'d  rnmi  clicriiiral  in-o.-.^ssrs  (.(•.•iiiriiiir  ii 
tlic  tissii.s  altlmni:!!  sum.'  ol'  tli.'  .-ivatiiu'  ami  ciraliniiic  incscnt  in  tli- 
fiiiMJ  may  appeal-  as  cicatinc  in  tlic  iirim'. 

Essential  Chemical  Pacts 

M.fuic    «<■    pr,M-c,..l    riirtli.T   Mitli    a    .lis,Missi,,n    of  tin-    mctalH.lism    ..i 
tii.'sc  impoitant  suiiMances.  if  uill  I„.  ncc-ssaiy  (..  vcf.T  In-icllv  f.,  sd.ii 
points   in    their  .•iicniistiy.      The   simpler  „{  tiie    t„„   l,n,|i,.s   is   ereatin, 

ulii,-Ii   is  metiiyl-(.uani.iine-; tie  aei,i:  ereatinim"  is  its  aniiydri.l.  hem 

forme.i  fr.m  .-reatine  l.y  the  r.-mova!  ol'  a  inohviile  of  water,  so  tliat  tli 
MI,  L'-r.ni])s  iK.eome  Joined  L.-icther  in  tlie  same  way  as  tiiey  <lo  in  lli 
forniation  .,f  pef,tides  from  airiiim  aeids  (  pa-e  .Vlftl'.  Th.'  i'ehiti,>nsliii. 
are  illnstrati'd  in  tlie  follow  in;,'  formidas: 

(  Mietli\l  1 

'    \ 
/         CI  I  coon 
MI       <•  ,1.) 

\  (:,,.  Ii,'!!,  i,h 

\ 
f  uuniii.liiii' )      Ml 

(creatine) 

<||         \       I'll       CO 


i  ! 


I 


Mr -f  I 

\  i 

\  I 

\  I 

Mf 

(creatinine) 

It    shoilhl    ]»■   iinteil    thai    unanidiiie    is  eh.sely    related   to    men 

y 

^**     *'  '■  ••""'  "'••I'    "'"'ii  en-itiiiine  is  foiiiK'd   fi'om  creatine  a   rim 

\ir, 


Tin;    Ml'.TAIiOI.ISM    OK    l'H(>ri;|\ 


623 


fDiiiiatidii  (K-ciii-s,  <;iviii<r  whiit  iiiny  lie  rcirfiidcd  jis  an  imidazolo  deriva- 


tive   (s.'e    |ia;r«'    fUI4).     Creafiiie    is  also    lelate.l    t( 


iliamiiio  acids,  arfrinine,  siiiee  liotli  eoiifaiii  fjuanidiiie  radiel 


one   (pf   tlie    iiiipDrtaiit, 


OS, 


MI. 


'MI     ('  \  and   to  iiistidine  and  llie  |>iirines    i  see  piijre  (l:!4).  Imtli 


il'    \\\\ 


\ 
NIL 

i-li     emitain     the     imidazole     lintr.      'I'l 


lose      relationsliip     nliiell 


creatine  lieais  to  urea  is  illustr.iled  by  tile  fact  that  urea  is  formed 
when  ereatine  is  snlijeeted  to  the  aetion  of  lioilinj;  li.irinni  hydrate.  Wlien 
It  is  oxidizeil  hy  means  of  |)otassiiiin  i>ermaii<ranate.  urea  is  also  formed, 
the   remainder  of  the   moleenle,   more  or   less   intaet,   hein^'  sjilit    off  as 


iii'lhyl-amilioaeetie  aeid  (CW 


/ 
\ 


xir-nr, 


),   also  known   as  .•■ 


sareosine. 


f'OOlI 

The  conversion   of  en-atine   to   creatinine   coes   on    slowly    in   afpieous 

-oliitions.  l)ut   is  mneli  ; jerated  liy  heating'  with  aci<l.     Heated  in  an 

aiit.ielave  at  a  temperature  of  117  ( *.  for  thirty  minutes,  with  half  nor- 
mal hydrochloric  aeid.  the  creatine  sroes  over  almost  (pmnf itatively  info 
'•i-eatinine.  1|  will  he  noted  that  the  creatinine  rinir  is  p.artly  oxidized. 
This  renders  il  iinstahle,  so  that  creatinine  in  the  jneselice  of  alU.alies 
has  the  power  (d"  red\icini;  metallic  oxides.  Like  ^diieose  it  can  reduce 
alkaline  solutions  of  copper,  silver  and  niercnric  s.ilts:  it  also  rediieos 
picric  acid  in  weakly  alkaline  soh.tion  to  pieramic  acid,  which,  hein<r  I't'd. 

I'uriiishes   us   with   .a   solution    the   stien-th    of  which    can    1 stimated 

cnlorimetrically. 

Quantitative  Estimation.-    .\ltl udi  the  i.reseii<-e  of  ciratinine  in  the 

(iiine  Ii.as  hecn  known  for  ni,-iny  \ears.  there  heiiin-  f,-,„u  |  (,,  •_>  ir,.j„„s  of 
It  in  the  Iweiily four  hour  urine,  little  prooievs  was  nuide  in  the  study 
■■r  its  mctaholism  hecjilise  c:f  the  ahseuce  of  a  leliiilile  method  for  its 
'■^limatioii.  Tin'  elalioiatii  ii  hy  l-'olin  of  a  coloiimet  ric  ipiaittitative 
method  f,ir  rn  hIIiiIih  .  dependiiiir  on  the  reduction  of  i)iciic  ;ii-id,  liiis 
i'liriiisheil  the  sl.iitiiio-  ,„,iiit  f,,r  the  modern  work  whi<di  has  lieen  done. 
To  estimate  the   innliii,    \,\    this    method,    it    is   usual    to    procecl    as    fol- 

'ows:     The  crcjitiniiie  < tent  is  first  of  all  determin.-d.  anofiicr  |)orti(Ui 

iif  urine  lieinj;  H,,.,,  heated   with   acid   in   the  autoclave  until   all   of   its 

'■'■''■■''i""'  •'■•i^  ' 'I  ' v.'rte(i  into  cre.-itiniih'.      .\  se<-oiid  -l<'teriiiination  of 

'■rejitinine  is  tiieii  made,  and  the  ililVereni-e  ln't  vv  ,..,.n  ih,.  uvu  is  i-alculated 
■  1^  creatilio. 


IPP 

Wi 

Ti  :  ^ 

w^ 

'I 

r    5 

F 

:i 

]■     8 

*    1 

I 

r! 

i 
1 

i|"    r 


l|. 


^f 


»■ 


1 


^1  ;! 


(124 


.mi;t.m«)i.ism 


It  sliouM  l,c  poiiiti'il  ..lit  that,  siii.v  111.,  rivatino  is  estimated  l.y  „„ 
mdireet  iiietlio.l,  tlieie  are  e.,iisi.lorai.Ie  ehniiccs  for  iiiaeeiiraey.  In.ie.M 
it  has  h.>eu  sh..\\n  that  ern.rs  lua.v  have  l.een  iiieiirml  in  some  of  Vr 
reeent  u.iik  on  a.-.-ouiit  of  the  faet  that  when  aeet..aeetie  aei.l  is  pn's...,! 
Ill  the  iiriiie  it  prevents  the  ereatiniiie  from  deveiopinsr  its  full  rediiein- 
I>ower  on  pierie  a.-i.l  in  the  e..hl.  s..  that  when  siil,se.,iientlv  tlie  nrine  T. 
h.Nitcl  with  aeid  f.,r  the  imrpose  of  eonvertins  the  ereatine  into  e.vi.il 

"""'•  •''^'  ''•■^tnietion  of  i toaeerie  aeid  allows  the  rediiein^'  power  of  tli. 

ereatinine  to  devei,.,,  to  full  infiisity.  It  is  obvious  tliat  this  would  n^ili- 
It  appear  as  if  .-reatiii.'  ha. I  he.'n  eonverted  into  ereatinine.  It  is  pai- 
tieiilarly  in  the  urine  ..f  .liahetie  patients,  in  whi.di  ae<.t,.aeelic  aeid  k 
present  that  niisfakes  are  likelv  to  he  nia.le. 


Metabolism 

When  we  eonio  to  eonsLh-r  the  inet.il.olism  of  ereatine  and  ereatinin.. 
we  find  that  there  are  reiiiarkahly  few  faets  definitelv  known  eon.-ernin.- 
It.    The  average  am..unt  exereted  daily,  expresse.l  as  the  niin.her  of  milli- 
frrams    of   ..reafm.ne  in  twenty-four    h.,urs    j.er    kilojrram    l.odv    weh'ht 
IS  known  as  fl„    <nati„l„f  cn,!]in,„t   (Shatferi.  «     For  a  lean  person  Uii. 
IS  about  25  mjr. ;  f.,r  a  .-orpulent  person,  about  20  mg.,  the  difference  in 
•lieatmjf  that  nn.sele  mass,  an.l  n.)t  bo.ly  weisrht,  is  the  important  faetm 
d.-termiiiin-  the  eoelTieient.     Further  evi.l.Miee  that  this  relath.n.ship  e^ 
isfs  IS  furnish,..!  by  the  faet  that  in  the  museular  atrophies  ereatine  ..x- 
eretion  is  distin.-tly  beb.w  m.rmal.     It  must  be  the  ma.ss  of  the  miis.-l.. 
rather  than  their  aetivities  that  is  the  .letermininjj  faetor,  for  the  ereatin. 
exeretion  does  not  lieeom..  in.-rea.se.l  by  mu.seular  exercise. 

Influence  of  Pood,  Age,  and  Sex.--  Alth.m«h  ereatine  .and  .•r..atiniiie  a, 
endogenous   metabolites,   it   must   be   remembered    that,    under    ordinarv 
•lietetie  eon,l.ti,.ns,  a  part  of  eaeli  is  .ierive.l  fn.m  these  substances  piv^ 
ent  in  the  f..od.     It   is  imp.ntant   tlieref..re  to  onsider  the  e.iiidition. 
under  which  the  ereatine  an.l  ereatinine  in  the  f,.od  appear  in  the  urin. 
Regarding  ereatinine.  it   is  pretty  well  ..stabli.shed  that  practically  ai' 
that  IS  taken  with  the  fo.,.1  reappears  as  ereatinine  in  the  urine.    SlialV 
has,  for  exampl..,  siie.-ee,l.>.l  i,,  re.-overing  76  per  cent  of  iuRested  .mv;,- 
mine  in  the  urine  exeret..!  durin^r  1wenty-..ne  hours  ffdlowiuf?  the  i^ 
frcstion  of  0.7  prm.  .•reatinine. 

The  conditions  for  the  excretion  .)f  creatine  are  more  complex      It 
present  in  the  urine  of  ..hildren  in  eonsid,.rable  amount,  but  in  that  ■ 
adults  only  as  tra.-cs.     In  the  first  years  ,.f  life  the  .Teatine  in  b,.^ 
urine  may  am..unt  to  one-half  of  the  total  creatine  and  ereatinine    1 
■t   been,,,...   Kindr.ally    l,.ss   an.l    prn.ti..ally   .lisaj.p.-ars   at    ab.mt    s,.v. 


Tin;    MKTAnOI.ISM    of   I'ROTKIV 


625 


years  of  Jifie.  Girls,  on  tho  otlior  liand,  contimic  to  excrete  creatine  until 
aiiout  puheity.  after  wliich,  ;iltIioiiL']i  unlinarily  absent,  it  reappears  in 
the  urine  at  each  monthly  sexual  cycle,  and  is  present  during  pregnancy 
■ind  for  sonic  days  after  delivery.  Feeding  creatine  to  children  causes 
it  to  appear  in  the  urine,  accompanied  nsually  hy  a  slight  increase  in 
the  ci'catininc.  The  same  results  can  he  oliscrved  in  women  during  the 
monthly  periods,  when  as  much  as  0.1  gm.  may  he  pi'csent,  and  during 
pregnancy.  Cicaline  is  also  pi'csent  in  the  urine  of  most  if  not  all  of 
the  other  mammalia.  Some  of  these  facts  are  shown  in  the  following 
t^il.lc: 


CKFATl.SINKX 


CKK.XTI.VKX  KXCKKTKD 

IV  :;i  in;.  I'KixK 


11.. \> 


(iiiN 


(  - 

e.CL'.-. 

(I.OLM 

:; 

0.(I.j7 

0.022 

." 

0.11  J 

0.025 

s 

O.IC..', 

0.0 

n 

(1.1. -.7 

0.0 

1.-. 

)i,!'.7'^ 

0.0 

10 

11' 


(I.IHVI 

o.n:i2 

0.1, -.7 
0.117 
0.201 


0.00,") 
0.00.1 
O.Ofif. 
0.020 

(1.011  _ 

(  Fiiini   Mutlipws.i 


AVhen  cicatine  is  given  to  an  animal  that  has  been  kept  in  a  starved 
conditioti.  most  of  it  seems  to  disaiM'car.  It  can  not  be  recovered  in  the 
urine  either  as  creatine  or  as  any  other  iiitrogenoiis  metabolite.  Tt  seems 
to  fnnctionate  more  iis  a  food  than  as  ;i  useless  substance.  The  possi- 
bility that  some  of  it  can  be  dcstroye(l  by  the  intestinal  bacteria  being 
;idmitted.  there  is  nevertlii'lcss  some  justification  for  the  view  that  the 
creatine  finds  a  useful  function  in  the  anabolic  process  of  the  muscles. 

Influence  of  Complete  and  Partial  Starvation.  -Althouu'h,  as  we  have 
sc(>n,  the  creatinine  excretion  remains  const;uit  when  the  anuuuit  of  pro- 
tein in  the  diet  is  irrcatly  i'edue(>d.  yet  it  does  not  remain  constant  during 
(•omi)lete  fasting  or  when  caibobydrafes  ;ire  entirely  withheld  from  the 
diet.  In  fastiiiij  it  has  been  found  that  ert.atine  appears  in  i>lace  of  the 
creatinine  which  has  disMi)|)careil,  so  that  if  both  creatine  and  creatinine 
are  determined,  xcry  little  if  any  diminution  will  be  found  to  have  oc- 
curred. Fasting,  therefore,  causes  the  adult  creatine  and  creatinine 
metabolism  to  become  like  the  .juvenile  metabolism.  As  pointed  out  by 
l^Fathews,  it  would  be  interesting  in  the  light  of  this  observation  to  see 
wliether  other  sub--tances,  ]>asse(l  in  the  urine  of  young  animals  but  ab- 
sent in  that  of  the  adult,  would  reajipear  in  the  urine  when  the  animals 
"'.vere  made  to  fa:;f.  lu  the  case  of  man,  fur  instance,  allantoin  would  be 
worth  investigating  in  this  I'cgard  ( p"fJ<'  •'-H''. 


H 


li   ',  I 


t  ,' 


^.i 


1  ij  Mi 


I! 

c  fi 


nil-'    < 


W 


fijtl 


Mi:TAIi()I.ISM 


■^   '"'"'■•"•  '■'■'''' ""'"'   "'■  -'""<■  "f  tl.o  .■.•..;.ti I,v  .•••...•,ti„P  ,■,,,,„.„ 

"''."";•"■'"'''>'''•"•'   i^  '■•""•'■'>■    ^vitl I,]    from    1l,..   .ri.t.    or   i„    ,Ii,,I„.t 

•■"""'"^-  "■'""■'■  '"  ""•  -'i^''^'- 'l-t-  .n,.lli.„s  in  ,„,„,  or  i„  ,1„.  .sp,., 


rtii 


";""'■■•  ""^i'l<'i-.-:M.'   IMH   ..r  tl,..  worl<   tl.Mt   l,,,s  l„.,.„  ,| „   „,„ 

I'l'.'-'  "I  tl,,.  snl,.i,.,.t  :,  MiHlinW  nf..Mi„w,lio„  w.s  nnpl.-v,.,!  wlnH.  .li,I  n..- 

'''^'' ;"""•"•"">■    '"•"  '"■'•'"".t    tl„.   iniln,., of  , t..,.-..ti,.   ,u-\,]    „„    ,1:, 

"■'■''•"" ^""""1"";  l"«t  ..v,M,   :,rt..r  .-illoui,,..'  i-.„.  il,is  ,,.,ssil,!,.  s„nr..,.  „■ 

"'7V,"''"  "•■'"  '"■  '""'""'"  """  ••-•■"!".'.•.,.,.,....  in  tlH.  unno.l,,.^ 
•■■■>yhuhy,h;ir.  .•,,...  i„„.n,|„.,iy  nirl,.,!,,,!!,..,].  if  ,.;„.|,oI,v,l,-.„os  „,•.•  ..i,„. 
to  .  ^t;,n,n^nMiMw,l,  f.,r  ,.x;,„„.l...  tl„.  .n...!,,..  is  ,v,,h 1  i„  i,s  urin.  I,, 

;;,'■••"""""•  ;''"'"""''  "'!^  "  i"  •"■•  '-"-•  "Iin,  ,.itlM.,.  prot.in  .„■  f.-.f  is  M 

IH'  unH.rnl  ..„n,.|„si„„  „l,i,.l,   „„,.  ,„.  ,,,,,„„    ,...„„  „,,,^^.  „I,s,.,.vati..Ms  i. 

I  Kit   .•M,^,n|,wh.Ml,.s  in  s„n„.  .;,y  nn.  ,v,ni>v.|    f.„.  ,!„.  p,.„p,.r  ,.o„v.Mxi„n 


Origin  of  Creatine  and  Creatinine 

Xntu  i,|,s,;„„]in.-  ,1„.  ,.n„n„Mt    nf  ,..,.„I1„„,    work   ,1,;,,    Iu,s   nvontlv  1,,.,., 
:'";"■  "'■  '1-  .-t,.,),olis,„  of  ........„iM,.  ,.„„,  ..,.„,„iMin,..  ..  knou-  vM-v  lit.l,' 

""'7;\'""^"'^"'~  " 'i-i"  ■■'•  'I'-  '""li.'s  in  tlH.  .MiMwil  orj..niM„       !• 

;:""'';'■  '"■""""^•^  • -"- "'-  p-i'i-  to  .„,•  ......t  o.,,.,,.  iM,t . 

t.;"-   ol    tlH.Hos,    „„po,.„n,    f,„.,s   so    n„.,.sl.l,lisl„.,l    M,;,V   Im.    of  int.MVst    MM,! 

"'  '•■''''"■      ""■  '"^'  •"•'•I'  i"  .•itl=...UiM.  sn.i,  ,.,  p,..,l,l..M,  is  fo  ,.o,np.-uv  ,1,. 

•m.oinits  p,.,.s,.nt   in  tl,..  v.-irious  o,...„s  ..,,,.1  tissues,  in  il,..  I,l.,.„i    ,„.,  ;„ 

-  --•'■<■<..    ;"'"-n..Moxi„K,„.M20„.„.,sofnv.,i„...n.lnv.,inin, 

""     ""^"" -n,.,.  ..h.H.  .  v,.,.y  1......  p, ,, is  i„  „„.  „„.s,.,... 

t-mn.^.   >,„.,„,,„   p,.,.,-.„t,-,..,..  ;,n,|   „,..  i„vo|„nt,-,ry   (inlosti,,,,!)   .nus,-!,.. 
••;•"  ••"mn.^  r,.]Mt,v..h    ,-,  s,„,-,ll  n,„o,„„   ,  Myn-s  ..,.„!   l-n,..,  -.     NVx,   to  f.. 

;'■''■"'     ""7'"^-     ^:"''     ■•'""- -    ' ti'-:      .1-     iovoiun.nrv      uu. 

'•'':^-   '■"""•""■  '"^"^  "'--"•      •••■'-   liv--.    |Mn.-,.,.„s.    ,l,y,.oi,l.   Ui.hM.^. 

.  th..  siii;ill..s|  ;ii,ioiiiit  of  iill  |„.in,j  f,„„|,l  ill  ,i„. 


''|>l>'''0.  i.tc.  .•ont.iiii  t 
M..0.1. 


In  .11  ti„...  ,,i.,.,.s  u-  n„.  ,|.,.  ,,,.„„„t  p,.,,,.„,,i f  „„.  ,.„,,,  ,.,,,..^„,,,, 

'•;■' -'t.-nn,.  ,.x,sK  ns  ,.,.,..,;,,,,„,,,,,   K,,,,Hly,l,..,.,.v...^^^ 

'"'";'':•■•""":;,'"  ""■  "'i' '■■•"i"i'^-  "1h.,v  p,-,„.ti..,.mv  ,,ii  is  ..v..,.,.,,.,, ,. 

-..„,„„.       ,|„.    ,s„  „,„„„,,,,,    ,.,,,,i,.nsl,ip,.,nv ,.„|,i ,..,,,„ 

'^""■''"" "'-'""l>.in.li,-at,.s,l„.„  ,1,0  ..,,,„ .„,.„,„' 


,.;,],•  ,,  .""•     "1    1  in-    III  IMC    IS   He 

""".""'"    ""■7-"' •.!..•    ti..u,.s.      T!,..    ,,H..,io„    is.    no,r.I,.,..    li,. 

"'"" '"'"   ' '•"'"/'"'".-'//»...'     S,...|,  .  t.n,sfonMn,ion   ,. 


1^ 


TUr:    Ml   TMiOI.ISM    OK    I-KIITI'IN 


627 


|)r()l)iil)ly  clTcctcil  Ity  iiijiiiy  dl'  llic  tissues  ni'  tin-  lioily  iilul  ccftiiiiily  Ity 
tliP  l)l()()il,  till"  ;icti\('  iiifciicy  ill  nil  ciiscs  licinir  no  dniiht  iili  ('liz.\iiii'.  That 
tin-  Mood  contiiiiis  siirli  an  cii/ynir  is  iiidicatctl  \iy  the  t'ai-t   tliaf  crcatino 

is  traiisfoi'iiicd   to   crcatiniin'   liy   lil 1   si'i'iini   iimrc  4|nickly   tlian    it    is 

wlipti  iiKM'dy  dissolved  in  water.  Kveii  lieateil  lilnod  serum  jiossessi'S 
some  of  this  jiower.  Tiie  liver  also  jiroltahly  Iniiiu-s  alioiit  the  traiisfor- 
iiintioTi,  as  has  lieeii  slio\\  n  liy  peifiisioii  exiieriuients,  and  liy  tlie  1'aet 
that  in  eases  of  i>iios))lionis  or  liydra/ine  pnismiiii!;  crealiiie  displaees 
ereatinine  in  the  urine. 

Tlie  ])roli|eiii  therefore  narrows  itself  down  to  tlie  (piestioii  of  llir 
iirifiiii  of  irniliiif.     In  tlie  li'.dit  of  elieiiiieal   knowhdtre  there  are  several 

|ireeiirsors  from  wliieh  creatine  miirlit   lie   fori 1.     One.   for  e\aiiii)le,  is 

ars'iniiie,  whieh  it  will  he  remeiiilpertvl  is  irnanidine-aiiiino-v  aleiianie  a<'id 
(see     pav'e    (ill.'i).       |*.y    oxiihitioii    this     iiiit;ht     In ;ne    eliallL'ecl     into    iruaiii- 

dine-aniino-aeetie  aeid,  whieh  liy  niethv  laiioii  would  then  he  idianu'id  into 
ereatiiie.  That  siiidi  a  |)roeess  of  iiii'th\  lafion  iiia>  actually  occur  in  the 
animal  liody  is  dednitcly  known,  for  it  happens  when  siudi  suhstances  as 
pyridine  or  naphthalene  are  iriven  with  the  food.  They  appear  in  the 
urine  as  nieth.xl  derivatives.  The  iiossihility  of  the  derixation  of  creatine 
from  aririiiine  is  not.  liowever,  liorne  out  liy  the  result  of  the  in.jectioii  of 
ariiinine.  for  siii-li  injection  does  not  ineiease  the  creatinine  in  the  urine. 
The  closely  reIat<Ml  sulistance.  ifuanidine-ai-etie  aeid.  when  fed  to  animals 
fraliliitsi  dues  cause  a  slii^dit  increase  in  the  e\c-retiou  of  creatine  fJalTet, 
aliil  also,  it  i>  said,  an  increase  in  the  creatine  content  of  the  muscle. 
Kveli  ill  this  case,  however,  hy  I'ar  the  Iari.'est  propcution  id'  the  admin- 
istered  triianidine-acetic  aeid  is  excreted   in   the  urine  unchaini't'il. 

The  larjic  ))ereeiita<:e  of  creatine  in  muscle  tissue  leads  one  to  expei-t 
that  some  relationship  must  exist  Iictween  muscular  metaliolism  and  the 
amount  of  creatine  ineseiit  either  as  such  in  the  muscles  or  as  creatinini' 
ill  the  urine.  [{eL'ardiin,'  the  latter  jxiint  it  is  definitely  cstalilishcd  that 
iiiilseiilar  exercise   leads   to   no    ini-rease   in    the   creatinine    exeretiou,    al- 

tlioiiuh   it   is  said   that   an  increas curs   followinu'  a   tonic  coutra -tion 

of  the  muscles.  With  reuard  to  the  creatinine  in  the  muscles,  no  definite 
results  indicatinir  that  muscular  metalinlj-m  idiaiiires  its  amount  are  on 
record.  In  the  liudit  of  the  tact  nlreacjy  htated  r(><.'ariliii^  the  i>rcsenee 
of  creatine  in  other  or^rans  than  the  iiiuscles.  it  seems  prohalile  that  the 
siilistaiic<'  has  rea!l.\    little  to  do  wiili  miisciilar  contraction  as  such,  hut 

rather  is  coiiccriii'd  in  some  wa.\    in  tlie  formative  iiietalioliMii  of  tin II. 

with  its  •reiu'ial  ijrouth  or  maintenance.  Indeed,  it  is  a  (piestion  whether 
creatine  is  an  actual  coiislitiienl  of  the  lixinir  tissue.  It  may  rather,  as 
lias  iieeli  sli-iiic.sled  !n  i'otlii.  lie  a  pn>|iiiiolcm  prndiicl.  represenf ed  diir- 
inij  life  1)\   creatinine. 


\u 


«  ■-.. 


r.2s 


METABOLISM 


•1  '         '  > 


(  .Tat.n."  appears  ,n  thr  urine  in  phosphorus  poisoniuR.  i„  enr.inoma  nf 
tlM-  iivor  an.l  .lunn-  postimrtnn,  involution  of  the  uterus.  It  is  not  <!.■ 
rived  from  the  disappearinc  uterine  n.usele,  however,  for  ereatinuria  als,, 
oc.-urs  after  eesarean  seetion  uith  removal  of  the  uterus.  (Veatin- 
ol.m.nat.on  is  not  an  index  of  eellulnr  destruetion.  for  it  has  been  fou.M 
arse  ...  a  .lo,.  in.jeete.l  uith  phh.rhi.in  an.l  maintained  in  eonstant  wei-^l,. 
.y  feedmv'  M.th  uashed  meat  (S.  I{.  Henediet).  Mu.seular  faticue  aK., 
loaves  the  ereatine  eontent  of  musele  unehansfed.  In  late  sta-es  o, 
nopln-,t.s.  ereatnune  a.-eumulates  in  the  i,lood  and  serves  as  an  index  ..f 
tlie  ^'ravity  of  tjie  eondition   (  pa-re  f..")|). 


■;!     J 


nl 


'1<      i    vh 


w  i 


CIIAPTKK  LXXI 

TIIK  MKTABOLISM  OF  PIJOTKIX  (Cmt'd) 

UNDETERMINED  NITROGEN  AND  DETOXICATION 
COMPOUNDS 

In  tho  pros-out  diaplor  uo  sliall  reft-r  briofly  to  tlie  gvonps  of  iiriiinry 
siil.stanocs  stylo.l  uiKlctoiiniiiod  nitrocpiiniis  oompouiuls  aiul  to  the  eoiii- 
pouiuls  tliat  are  oxorotod  in  tlic  urine  as  tlie  result  of  the  combination  in 
the  body  of  certain  toxic  bodies  Avith  ehemieal  sul)staiii'es  tbat  render 
them  harmless  fdetoxication  compounds). 

Undetermined  Nitrogen 

Included  nnder  nndetermined  nitro<;en  are  amino  acids,  pei)ti(les  and 
basic  substances.  The  amount  of  amino  acids  and  peptides  in  normal 
unnc  is  very  small  but  may  become  considerable  in  disease,  especially 
of  the  liver,  when  leucine  and  tyrosine  may  appear.  The  presence  of 
traces  of  amino  acid  and  i>c))toiie  in  normal  urine  is  to  be  expected, 
for  althoujrh  the  actual  concentration  of  amino  acids  in  the  blood  is 
never  very  Rreat.  a  certain  leakafic  of  amino  acids  nuist  occur  into  the 
urine. 

The  peptide  is  sometimes  known  as  nxuprofric  afid  It  becomes  dis- 
tinctly increased  in  phosphoriis  poisonins  and  in  such  conditions  as  are 
accompanied  by  excessive  protein  metabolism.  The  hnsic  comtiturtiix 
include  such  substances  as  trimethylMMiinc.  etliyhimine,  putrcscine  and 
cadaverinc  (pajre  .WiK  and  there  are  i.rohably  many  more  of  a  similar 
nature .^  :Many  of  these  substances  are  similar  to  tho  .so-called  ptomaines 
found  in  meat,  etc..  and  they  have  been  called  the  ptomaines  of  urine, 
from  which  they  can  be  isolated  by  renderinsr  the  urine  alkaline  and 
shakinp  out  Mith  ether.  It  is  probably  to  the  presence  of  these  sub- 
stances  that  urine  mainly  owes  its  'oxic  action. 

The  Detoxication  Compounds 

Certain  nocuous  substances  are  produced  in  the  intestine  durin?  the 
dipes-tive  process  rsee  paee  "lOn,  and  othei-s  may  result  from  the  niota- 
bolic  processes  in  the  tissues.  To  Ruard  against  the  harmful  action  of 
these  substances  on  the  organism,  they  become  detoxicated  in  various 

Cy29 


ill 


'■  ..  I 


m 


m 


I:  ;  if' 


r  1 


(>l    • 


'1   ■■  I 


rf: 


■,< 


.  i 

■  1 

H 

■  1       ' 

! 
j 

•t 

6.10 


.Ml  TA not. ISM 


III 


w;i\^.    Mi;iiiil>    l,y    rcniiinir   inrrt    .•.iiii|h>iiii.|s    uitli    ..tli,.,-   Mil,s|„iin-s,    |. 

''""'■"■'>     "i"'    '-il> II.    stil|.lniii,.    ;i.-M    „!•    silwuiciiii-   i,ri.!.      Tin-    n, 

|'"iiiiil  iliii^   I'lirni.'.l   is  tli.Mi  cN.-ivl.'.!  in  tlif  iiiiiic. 

Hippuric  Acid,     (il.v ,.,„-,. II   is  iis,..|    „|,.|inly   to  .|,.|,,xi,.,it,.  li,,.  I „,;. 

•'"■'''   "''''■''   '■'■^""■'   '■'■ 'I Ni'l.iti r  ll„.  nroiiiMti."  Mii.stiiii.Ts  piv. 

••Ill  ill  l.ii-,.  .|M.iiititirs  in  vr-.t.il.I..  r i  ,iM.|   fniit     |.,nli..,il„.lv  in  cr.n, 

I.nri.'.V       S,.|r„.    l,..M/,.lr    j,,.!,!    „I,IV    .-lis,,    |„.    |.,„,iM.-r,l    I,.     |j„.    l,V,..-,l<,lnU  , 
"I    ""•   ••nnniati,-   ui„ii|.  uf  tl,,.    ,„.,,|,.m,    iiMilrn.l..;    plinix  l,-i!;iniii.-.    fnr   .a 

.•lllipli..    -iws    lis,.    I.,    I„.n/ni,.    M.-iii    l.y    h„rhr:,l   .|rr,„„|„,si|  inn.       Til In 

'""""'    I''"'' '    '•<   l''fl""-ir   '/./•'/,   this   iMIiir    iii,|i..;,tii,..'   that    it    is    |.|Vs|.,lt    in 

l.-i.-.'  .|i|.-iiitili,.s   in    ||„.   iirii,,.   ,,f  th,.   Imis,..   .ms    it    is   .-ilsn   i„    tlir   nrii i 

.■ill  Iictliiviiniiis  aiiiiiials, 

llirpiliii-    .H-iij    is    l..iizn\l-.rlyi-iii..      I'll    .Cn.MI.CH.COdll^    all,]     it 

'"'"    ''"■■"'''>     '"'    I"'"' ■•'    ill    til-'    l.llHllalnrv    |,y    lH-i.|..'ill-    l.l-.'tll.-r'lHMI/uvl 

'•Iilipfiilc   M  ith    ul.viM-dll.    Iliiis: 

'"<••'    <1     II     IIV.CII  conn     <'IM().\l|.(||((M»ir     MCI 

ilM.ii/nU  ..l,l„,i,|,.,       ,,_.i,., in  (Iii|.|.uri.-'a.M.n 

I-.i.l.T   .ir-liiiaiy    ili.t.ti..    n.n.litinns    only    a    tn f    I,!,,,,,,,-!,.    ..,.-i,I    i. 

IMvs.Mit  III  til,,  ui-ii r  ,„,.,„,  |„|,  , 1,  |.,,j,,,,.  ,|„,„„i,i,,^   2  -riaiMs  a  .hn 

'"'"■  '"^'ii'il'l''-   'iiiiv  apprar  wh,.,,   tlir  .liH tains  a    lar-r  i>n.i),.iti.m  ,il 

fniit  nr  v,.-,.tahl..s.     It  is  ii,it  l<ii„u  n  to  iiii,I..,,.o  any  .•harartcristi,.  vaiia 

tioiis  ill  ilJM.as...     Th,'  hcn/ni,.  a,M,I  «  liirli  is  itai 1  in  cTlain  raniid 

'' ^"^  ■'■'   I'l'-'iv.itiv   also  ,-oniI,iii,.s   in   tho   ]u»\y   with    -lyocoll.   s,.   that 

any  t,.Ni.-  HT-M't  «  hirl,  it  mi-lit   pro,!,,,-,,  is  prarti.'ally  n,.-li-.'il.l.',     Tli,i. 
i-  crtaiiily   i„,  v,iy  ,.vi,l..|it    ivas,,,,   why  ,.ann.-,l    foo,ls  containiii!.'  U'uy.ul- 

•■'"'''  '^'""''''  '■'■  '••'' '•''•  I'll-  ill  -^1  I'.-ii'  iis  th,.  U'yy.sH-  a.-i.l  is  ..on.M.rn.-.l.  tll..^ 

'■'III   !"■  11"  i.'i"i'''  t.iv.i..   than   a   .li.'t   .-onuMis-.l   lar-,.'v  ,.f  voirrtahh's  an, I 
rniit. 

Tliis  ,l,toNi,.;ilion   ol'  !,..„/,,,!,.  a,M,]    iv.|iiiivs  th..  piv<,.n.-c  in   tin-  (n-an 
'""     "'■     •■'     •■"ll^'.llit      supply     .,f     .j|y,.,>,M,||.     whi,.),.     it      will     1„.     ,.,.,.;,l|,..l 

IS    tl,,.    lov,,.st    in    th,.    s,.,i,.s    ,,r    aiiiin,.    .....hls.    Inin.,'    aniinoac(.ti,.    a..i.i 

ifir.MlrtHlII      .  1,      is     p,.,..,.„t      i,|      ,_r,,..|t,,.,,      ;,„„„,„,      j,|      ,|„,     ,„,,,,,;„     „f     ,|,„ 

.•o,in,.,.t,v,.   tiss,i,.s.      It    is  sai,l.   how,.v,.r.   that    not   nmn.   than    from   '_'  t.. 
:!...  11,. !■  .•,.nt   ,.r  i;ly,.,„.,,ll    is  a\ailal.|(.   in   tli.-   i>n>t,.iiis  of  tlio  liody.      .\I 

"'""-''   "''-  •■ii""iiiit   of  .Jy,., II   wonhl   amply  snni.-o  to  .l,.toNi,.at<.  tl,. 

'"'"^"'''  ■•"■'''    I"'"' 'I   I'y   »!.,.  niHaholisiM   ot'  th..   loo,l    in   ..arnivora.   i' 

IS  .piit,.   iiia.l,..iiiat..    Tor  tlws   pnrp,.s(.    in    the  .-as..  .,f  )i,.rl,ivora.   an.l    th. 
.|ii..sti,m    naliiially    i.i-s,.nts    ils,.|f   as    t..    uh.-iv    th,-    .,Mv,-o,.oll    in    tli,... 
.■I'liiiials  ,.oni,.s  In. III.      It   is  sai.j,  for  ..xaiiipl...  that  .,f  th..  t..tal   nitn-u,)- 
-■.M-n-f;oii   in   liMi.!;,,,,,  .',(1  p.r  ..,.nt   may  appi.ar  as  !.|vco..oll   un.l.T  ....i 
tain   .■on.litions.     Tins,,   l-i.-ts   in.li..j,t..   that    thr  ..r-anism    is  capal.l.^   oi 


nil;   Ml  lAiiiii.isM 


III'  nmriiN 


tl.'tl 


liKiiliii'iiii,'  iif"   }rl\ccir.ill  I'.ir  iiMir,  jinil  it  i>  inlcic-'tiiit.'  t iisi.l.T  Ikiw 

lliis  i;l>ci.c<il|  niiiy  hi'  (|cri\  cl.  A  very  iirolwililc  smiiff  is  \i\  Miillii'sis 
lii'twrcM  ;Miiiiic.iii,i  iiiicl  ;.'l\ip\ylif  iici'l  iCllO.  COOII..  'I'liiil  !.'iy<ixylic  iicitl 
or  its  iilili'liydr,  uly..\;il,  is  ii;i.|il\  |>i ihIiii-i'iI  diiriim  iiirt;ili..liMii  Irnni  chc- 
lHiliy(Il;ilc,s   jilhl    lliiit    .iiMliioliiii    is   iilwjiys   ii\  iiililMr    wrnilil    scciii    t(i    Icinl 


Slim.'     SII|l|M.rt      to     I  hi,      \ir\V       ISC..      |M..r,.     I'lll, 

iVdiii   irly(i\:il  mill  iiiiiniiiiiia  (n-ciiis  tliiis: 


'I'lir    s\  iitliisis    III'    •r]\  i'i>ri,|| 


M.riii'lH)      Ml         (11  Ml  ((Mill. 


J^..\.■lh 


:lv '.Ill 


Till-   liliKiii!.'   up   III'   •.'lynii'iill    willi    licii/iiic   iiciil 


iMTiirs   ill    till'    Kidiirv. 


II  till-  kidiiry  is  iciiMiM'd  I'miii  ill..  i-iiTiil;iti.iM  in  ili..  m.-ijiiritx   nf 


llijit   piiidiii.(.  Iiippinii-  ;ici.|  ill  iii 


lion     no  liippiirii-  ..icid  will  ; iiiiiiiinti 


i''-'t'  ,.|iniiiiiit      III..  I'lililiit   iM'in.,'  :iii  rxiTp 


an   isol;it(.ii   p(.ii"iis..d   prcparalioii   ,,\'  ilir   liidiicy   pmd 

provided  |p..ii/.iic  ill-id  is  addrd  lo  tlic  pnfiisi.ni  lliiid.  mid  iii|.  l.it! 


ill  till.  Iilood.     <  )ii  till'  otiii'i'  li.'ind, 
iH'i's  liippiiiic   ai'id 


IT  also 


Viri'll,    wlilrli    is    lust 


rrlinil    l)\     llsiiii;    i||. 


liriai    sniiiii    (  |,.ick..'s   s.i] 


II- 


contaiiis  an   .•iliiindani..    oT  ii\ 

lihrinatid    aiti'iiali/t.d    liloinl    insliad    oi'   juii 

lion).     Tlif  iii.|.|.ssity   111';!   piciitiriil   siipplv    111'  o\y"j..|i    is   riiillic|.  slmw  n 

li.v  (lie   t'ai't   tliiit.   il'  lli(.  liriiionliitiin   of  tin'  lijond   is   iciidcrcil    incapalili' 

of  cairyiiiy:  (>.  I.y  liuliMin.,'  i-iirliiiii  iiiiiniixii!(.  ._'as  lliiiHiiili  it.  no  syiitJK.- 

sis  of  hippiiiic  ;ii-ii|   will    I-,. Milt    finni    p..rfiisiii._'  tlii.    M I    llir..iii;li    tli.' 

liidiii'v.     Tilt,  actual  dicniical  pr ss  liy  wliidi  llic  s\ntlicsis  occurs  (df- 

liydratioin    is  siinilMr  to  tliat   liy   wliicli   p.ijypcptidcs  arc   fornit.d  liy  tlic 
union  of  amino  acids,  nr  creatinine   from  creatine. 


M'.ir  CO  utM    II'  ii.\ci[.('ooiii. 


(ilv 


cocoii  ma\    III 


d,   particiilarlx    cl 


Used   fur  deloxicalii.;,'  otiier  siilistanees  tliau  Ix.ii/oic 
loiic   acid,    foriiiint;   the   til itrni-I'dHi-   uriil   of   llie    Iiile 


i/o;c     aciij     II... 


(.s..e    pae:..  4!U)    and    pheiiyh|..i  t  ic  acid.      In    jiiids     the     Im.i 
comes  comliined  with  iliaiiiino-v  jilcrianic  acid  or  ori.itliiiie   iNII.     ("If. 

<'H,, -('IF.  CM     Ml,     (■(Mi||i   ill  phice  of  .,dy 11,  so  that  in  the  mine 

of  these  animals  in   plai f  hippinic  acid  a  emiipouiid  called  (,niilh uric 

iiciil  occurs. 

It  is  of  importaiH-e  to  poinl  out  here  that  this  |iairinir  of  aromatic  toxic 
substances  with  <-(.rtain  of  Ih.-  mi'taliolic  products  of  the  oruMiiism  has 
fre(|iienlly  heeii  found  an  excellent  (.xpcriiiieiital  method  for  demon- 
stratiiiir  the   i)resence   of  inlermediarv    m(.|aliolic   sul'staiices  tliat    otlier- 


\\\ 


se    would    not    hav(.    appeared    in    the   cxci-eta.      Tli.'se    siilist 


thus  div(.rted    from   tlu.ir   normal   con 


rse    III    nil 


inces   are 
taliolism    so   as   to    foiiii 


neutralization  or  detoxication  comiioiiiids.    (dycnronie  ai-id 


is  an  exain])le. 


■p 


■f      } 

r 


iii 


'  :    Jf 


I 


i'li  '1 


'  \    ■-! 


r 


1 

i 

f 

4     ' 

j 

■; 

li 


8» 


MKTAB0IJ8M 


Ethereal  Sulphatei  and  Olycuronatei.  Tl.c  other  substances  use.l  for 
•letoxicntion  purposes  luc  sulpl.urii-  nnd  Blycun.nic  ai-ids.  I'l.c.ol  „„.| 
Its  dmvHtive  <MTHol.  aft.-r  L.-in^  al.s..rt.rd  from  the  i.itestinr.  in  fl,, 
.•ontents  of  whi,-!,  th.-y  are  ,,ro.lu,Td  l.y  the  ha.-torial  .lecompositio,,  ,„ 

'"■""'"'    '"""   l"'^''•   '""I    '"■<■'" i"'-.l    in   th.-   IhmIv,   pn.hid.lv   i.,   tl,. 

l.vcr.  Av.th  sulphunV  a.-id  ,„■  with  u'lyuronie  acid  to  form  the  sulphat. 
..r  Klycuronat-.  Th."  aromali.-  sulphate  further  e..mhinos  with  potassiu.,, 
t..  torn,  the  s,M.an..d  ,/A,n„/  s„l,,h„l,..  as  whieh  the  suhstanee  is  eNen,,,! 
in  the  nnne.  A  small  amount  of  phenol  mav  however  appear  in  tl,. 
unne  un.-hanjre.l.  As  we  have  alrea.ly  seen,  the  sourees  of  the  phe,,.' 
in  the  intestm,.  are  tyrosine  and  j.henylalanine  (s^e  pape  UWiU  an.l 
sinee  these  amino  neids  are  also  present  in  the  tissues,  it  mipht  he  su,. 
posed  that  some  of  the  phenol  sulphate  of  potassium  present  in  tl,. 
iir.ne  eould  eome  from  the  tissues,  It  is  usually  assumed  that,  howver. 
derivation  from  the  tissues  does  not  oeeur. 

An(vther  ethereal  sulphate  is  hulo.nil  ,„lpha>r  of  pntn.sium.  whieh  re 
suits  from  the  absorption  into  the  Mood  of  the  ind..le  nnd  skafole  pr„ 
dueed    by    u.testinal    putrefa,.tion    from    tryptophane    (see    ,.aire    50", 
Immediately  after  absor,)tion  indole  is  oxidized  to  indoxvl.  whieh  th"-, 
eombines  with  sulphurie  aeid  and  with  potassium  to  form  indoxvl  sn' 
phate  of  potassium,  whieh   is  the  well-known   imUmn  of  the  urine       \. 
Ml  the  ease  of  phenol  sulphate  of  potassium,  none  of  the  urinary  indiean 
seems  to  eome  from  the  normal  metabolism  fof  the  trvi.tophane^  of  tl,. 
tissue  proteins.     It  is  a  mneh  more  reliable   indieator"  than   phenol  s„l 
phate  of  potassium  of  the  extent  of  intestinal  putrefaetion    but  it  aN^ 
beeomes   increased   in   ammint  durinsr  putrefaction    in   the   bodv  itsell' 
as  'or  example  in  abscess  formation. 

The  amount  of  indican  in  the  nrine  may  be  rouphlv  £rau,rcd  bv  oxi 

clmns  the  urine  by  means  of  hypochlorite  nnd  then  shaking  out  will, 

chloroform.     If  the  resulting  extract   is  more  than   li.ht  blue   in  color 

It   indicates  excessive   putrefaction.     A   negative   test   does   not   neces 

warily  mean -that  intestinal  putmfaetion  is  absent,  but  a  marked  positiv, 

ost  a  wa.vs  „Klu.ates  that  it  is  oceurrin...     Skatole.  the  methvl  deriva 

ne  of  .ndole,  may  nndcrpo  similar  processes  and  appear  in  "the  urin. 

mn^  excessne  intestinal  putrefaction.    Its  presence  in  the  blood  some 

tines  confers  on  the  breath  a  distinct  fecal  odor,  for  this  bodv.  as  i,. 

name  indicates.  ,s  that  to  which  the  o<lor  of  the  feces  is  due      " 

is  ^Z'Z  ""')  *''t    '''"  "''''"""^  "•"'^   ^"•-  -^^t-'-tion   processes 
s  of  the  nature  of  a  dextrose  molecule  with  the  one  end-^ronp  oxidi.cl 
to  earboxyl  rCIIO  -  (CUOm,  -  COOII).    It  is  probably  produced   ,nde 

,z  ir^r' "''^^^''"'■^"^ '" '''  ^"'"^'^^  '.ody.'bJttdest;::!:! 

i.nless  «hen  .such  poisonous  substances  as  camphor,  chloral  hydrate  or 


dVl 


THE    MHTABOMHM    Of   PHOTKIN 


63.1 


rcrtnin  nroinafic  nlci.holK  me  tfivi'ii,  when  it  \h  used  for  the  purpuse  ul 
(letoxipntiiiK  thorn.  The  rcsultiiiK  Kl.vt'iiroiuiti-s  hnvo  rcdueiiiR  powerft 
and  may  be  confnse<l  witli  (jlueose  when  prcsont  in  hiiKP  nmount.  Oly- 
I'uronntps  may  lie  flistinirnishod  from  tflucose  in  the  urine  (1)  beeauBe 
they  are  levorotatory.  and  (2)  lieeausc  tliey  (hi  not  ftrnient.  The  free 
acid  itself,  however,  is  dextrorotatory. 


i   t 

i    : 


li   ! 


I         ft 


^.4 


If 


r 


i': 


I 


'I''' 


''( 


••.i-  if     ^ 

[A    ' 

111 


,1 


3! 


<n.\iTi:i{  i,x\ii 

I  l.'h    ACID  AM)  TIIK  I'l  i.'im:  M(»i>ii:s 

Introductory,     Th-  |.,nii.M|..-,i I.v  l,ii.|,|.v   ir.-.i,....!  ..rijmii,.  rl,..,,,!.. 

"'  ""■  '"^— "f'-ili r  l,in,.|M.iMi.-iil  pr-Mrms  Ims  I.i»m..'1iI  ohv  k.i.iu  I,..|.., 

"f  th.-  liiNtniy  of  111.-  i.iiniM'  .sul,s|„„r..s  in  )!..■  iiniin.-il  ImmIv  (V..m  n  st,it, 

"•"  '■'"'"''  '■""'    t''"'^'*""'^    I )•  s>st,.,„    .I,,,!    sri.Mitilir   il.-rufil.-v       Tl,. 

|..'c-iihar  s„h,l.ilily  ivarti..„s  „(  „■!.•  ii.-i.l  ami   its  salts  an.l  ll...  .lisn.v..,  ^ 

of  lilatrs  ,n  jTMitv  .Irposits  srr       |  f.  inaKr  >„!.•  ari.l  ....■tahnlism 

til..  ...irlirst    ivsranli    pn.M.nis   i„    |„,f|,    ,|,..   n,,,,,   ,.,i„i,.   „„,,    „^,.   ,,,, 

.•li.-mM-al   lalM.ial-.r.v.   I.iit    tl,,.   rarlirr   r..sults   wiv   ,.rarti,.allv    val,.,.!,>. 
'"""•    '"■'••"""■  ""•>   '"■'■'■  '"■ "•••"-  .-ll"!  I'.irll,   „s,.  |li..ir  iii't,.r,„vi;,ii,. 

"•■'^  ""'•".'•^ '"  ""■  ^''"'■'"■"'  "'■  ••^•"  ""•  i""-t  .•lrMi..nlar,v  fa,-|s  nmrrnii,, 

till'  fliriiiistry  (if  iii'ir  ac-ii|. 

Mrfof,.  aii.v  r.-al  pincirss  was  |...ssih|,.,  a  ,.l,.aii  sw..,.p  lia.l  I,,  lie  tn,nl. 

nf  all  111..  .,1,1  s|>,.,.Mhiti..iis  an.l  liyp..tli.'s..s,  s.i.-li  as  lluit  <lii.'iiili(..|  l,y  i|„ 

l.i«li  s,„ii,.li„^,   nan f  "  i„-i,..a.-i.|  .lialli.sis,   '   an.l  a    f..mMlati..t,   ,if  „ 

•■iwat..  .•li..|ni..al  kiiou  l,..|.,r..  ..stal,lisli,..l.     This  f,Mni.lati<.ii  is  n.,u    «,.,„|.  ■ 

fiill.v    .•.,nipl..t.'.   an.l    a   supcistiiK-tiiir   ..f   l.i.,..li,.i,ii..a|     fa,.t     is     alr.a.l 

l..-jriniii.i>r  1..  m-„u    .ip.,ii   it.      In   tl,,-   pn'snU   .-Iiapt.T   «.■   sliail   ..xami,,. 

■'*"""'  "'■  ""•  "'"^'  '"i|""taiit  tril.uti..iis  tliat  liavc  nii..lc  fliis  pi„iriv^ 

p,issililc 

As  in  til,.  stn.Iy  ..fany  ..th..r  ptoM,.,,!  ..f  in..tal...lisni,  w.-  nnist,  l,.,«..\,.i 
nial<..  ,.ni.s.iv..s  familiar  uitli   tli,.  main   fa.-ts  .•,.n..(.niin,_'  tli.'  du.misl i 

nf  til,,  pniin..  I„„li..s  aii.l  of  tli..  tissn i.stitu.'nts  into  tli.'  comp-isiti,. 

..f  ulii,-li   tli,.y  .Mt.r.   iM.f.n-..   in-o..,...,|ii,sr  to  tli..   nior.-  stii.-tly   l.i.,l,.i;i,... 
asp.'ct  .if  tlif  Mili.j.'ct. 

The  Chemical  Nature  of  the  Purines 

Hy  .•II.  ..Naminati.in  of  tl nipiri.-  f,.,- las  ,.f  tlic  pniin.-s  .if  l,i.i..l„.M, 

"■•'•'   i'i'"''''^>-  it  "ill  1, s..rv(.,l  that   th..y  ar.-  all  .l....i\  ativ.'s  .if  a  siil 

Stan.-.,  purl,,,.,  „jii,.|,  allhoi.ifh  in  its..lf  ,if  ,i portan.-o  is  intcn-sti,,.. 

'"' ''  '•'''■^'•^  "^  ""■  ''■■'>.''•  Mil.sian.-o  fr.im  which  th,.  othin-s  nw  .l..|-i\,.  ,: 

The  list  is  as  f.iil.iws: 


I'liriii,.         .         .  (Ml  x^ 

iryp.iNiintliint.      .  <'.II,\,'o  .M,mc,x.v|.iiiin,.        i 

■  ''UN, Ml,  Airiiim  |,ii,iii<.  I'liiini' 

•  •'.MN.O.NII,  Aiiiiriii  ii.\v|,iiiiii,.   I 

•  *'i".^."a  TiiiixypiiiiiK. 


A, I*. nil!,' 
Xiintliinc     . 
<Iii:,iiiii.. 
I'ri..  !i,-i.l     . 


I  Kir    Ar||»    AMI  riiK    fl  KIN>:    II11HIK..S 


«.'l.' 


Till'   (iiHt    ii\ii|jitii>n    |ir<M|iii't    (>r   imriiii'    is   li_v|M>xiuilliinc,    wliicli    I 


WIS 


Inriir  li' 


iri'ii  known  IIS  II  I'liiislitiD'iit  i>r  nn-nt  cxtiin-t.     Ailcnini'.  the  iimino 

ili'iiMiliM'  ol'  li.\  |iii\,iiitliiiit' nr>  in  i-ntiil>iniiliiin  \\i\\\  titli<>i'  HiiliHtnncrs 

in  till-  iiiicl.'.if  iMiilfiijil,     Tlic  M ikI  i>\ii|iitii>ii  pin.liirt   in  \iiiitliiin'  nii'l 


its  iiiiiini)  <|i'il\ii)i\r,  irniiiiiiM'.     'I'liry  mciir  in  tl 

Minlliiiii-  iiikI  inlrnini'.     TIh'  Itiiflii'sl  iisiiliiiiiiii  |ii-.m1iicI  nf  nil  Ih  tli 


ir    <illil<'   |ili'liM's   iis 


liypi 


i-ll- 

l>ni>\Mi  iiriiiiii-v  ciinHtitii.-nt,  uric  ihm.I,  ulijrti  nuiy  tlicicfnrc  l.r  clictnicnlly 
ili'siurniili'ii  (IS  tfii>\\  pill  inc.  In  ii<|i|itiiiii  I..  Ilic  purines  of  iininiiil  oriirin, 
ilnrc  !iic  also  n  rliiin  ones  of  M'urfnlpir  orii;in  I  he  mctliyl  purines,  wliii-h 
i\isl  MS  llir  iilkiiloiils  i<(  till  iiml  I'oll'cc  n.iMiclv.  cjilTfinc  tln'oi)i'oiiiiiio, 
<l  II 


Mill    MM'IMiV 


To   iinili-rstiiiiil    ll Iii'iiiirtil    siriirii 


IT   111"  tliis   irrmip   of  siilistiitn-cs, 


it    is    pcrluips   simplest    to   stiirt    with    tluit    <>f    111  ie    iieitl.      Tills   coiisists 
essentially  of  two  urea  nioleeiili's  linkeij  lo^etlier  liy  a  ci-ntraj  eliaiii  of 

'•"■ arhon  atoms,  as  will  lie  exiiient  from  the  afcompiiiiN  jn^f  .siructurni 

lormiila: 

II V    CO 


I       I 
<H'    (•     Ml 


\ 


/ 


I      I' 
US'   (•    Nil 

fiirc:i)       ( 

I 
(I'l'tit  i;il  ctiiiin ) 


men) 


This  striietiire  can  he  shown  l.y  methoiis  lioth  of  decomposition  nnd 
of  synthesis.  When  iirie  acid  is  decomposed  hy  oxidiziiii;  it  with  nitrjp 
acid,  it  yields  urea  and  a  residue  ealleil  alloxan;  or  it  can  he  synMicsizcd 
Iroiii  urea  and  trichlorlaclamide,  a  derivative  of  lactic  acid,  which  it 
will  he  rcnieiiiliered  contains  three  carlxui  atoms.  Tile  changes  involvcii 
ill  lliis  synthesis  will  l>c  maile  clear  liy  examination  of  the  accompanyini,' 
striietiiiiil    formula,    in    which    the    manner    of    luoduction    of    the    hv. 


products  of  the  react  iiui  (  Ml  ,  II ,( »  aiiijl  |( '!  i 


are  shown  liv  dotted  liiu 


NH.    !  II   Ml        (•  ~o 


/ 


/ 

/ 

;  ii"i 

ro 

\ 

:  CI  \ 

\ 

\ 

\ 

(lllCil) 

Ml. 

11  n 

II  !   c       (III  II  :    Mt 


\ 

C( 


<■-     ;    CI    It     :     Ml      (urea) 


(trirlilorlartamldc) 


i 


'  ■  1 ,  ,  1 

■     ■  1  ii 

1       '\ 

^  .13  I  1 

J:' ^ 


U\\\ 


f  .1 


ili 


T 


■,j  nil 


I    ^ 


t 

1  'i 
^1      ) 


C36 


METABOr.ISM 


By   miUlor  oxi.Iation   bv   moans  of  potassium  permanganate   in   the 
cold,  une  aeul  becomes  quantitatively  converted  to  allantoin: 

C,ir.X,0,    11,0     O    (\H,\/y,  +  CO,. 

(uric  acid)  (allantoin) 

The  importance  of  this  transformation  lies  in  the  fact  that  in  m,,^t 
annuals,  n.an  and  the  hi-dicr  apes  being  exceptions,  uric  acid  is  thus 
decomposed  .„  ,]„.  animal  body.  The  structural  forn.ulas  for  the  otlnT 
punne  bod.cs  m  relationship  uith  those  of  purine  and  urie  aeid  are  j,mv,.„ 

Purine  itself  has  the  follouinir  slni<Mural  formula: 

II-     ('•      (•       Ml; 

I!     II        \ 

Ii     ll  •"■■II 

'      'I        // 

^^'"''  ""'"' '■  '''-'ir'-"  'I-  M s  i„   ,„„.i,„.  :„-,.  „„,„1„ I  „s  s!,„«„., 


n\-c=o 

II      (•     (•     Ml 


I        I 
O     (•      C-Mf 


\ 


(l.vp...x:u,tl,in,.,        rn„.,,„„.l„r)  f  x:o„l,in.-)  f  2  (Icxv,,,,,  in.) 

N     <•   Ml 

I  I    '  "r*: '» 

"-f^    <'-Mi  iix.r    ('-Mr 

\  II     !!         \ 


■  I     11 


<•      II 


// 


X-C-     N 

ri'li'iiiiic)      iC,  ;irMinu  imriiicM 

IIX-  CO 

:        I 

OC      C  -  NH 


('   II 

I'      !!  // 

X-C-    X 

u'lKiiiiiic)       (L'  riiiiinci  Ii  iix,\  piij  in. 


!         11  CO 

I     ''      / 

iix-  r  -Ml 

T'ric  arid    i ::  iV^  I  rinxyimriiiPi 

The  substances  with   wbi..h  the  purine  bases  are  most  closely  relat.M 
are  the  /jnn.nidine  hasrs.     Three  of  these  are  known: 

thv.nino      rMI-ro  ..v,o.ino^    X^C-Mf,         .,„|    ,„«,.i|     ^XM-C, 

'^O     ecu,  rn    ru  ^l,     ^„ 


Ml       "11        ); 


XII      (11,; 


xir-rii). 


IKIC    ACID   AND   Tin:    I'KRI.VK    BODIES 


637 


From  an  pxiiniiiiatioii  nf  tlip  stiuctiiral  formulas,  it  will  be  seen  that 
they  are  more  or  loss  rolatcd  to  purine  (lia\ ;  (?  one  of  the  urea  radicles 
omitted),  althou<rh  it  can  seareely  l)e  douliti'd  that  ihey  exist  as  separate 
constituents  of  the  nucleic  acid  },M'<)up  in  the  animal  body,  and  are  not 
ilci'ivcd  from  purine.     Tliey  arc  jirimary  products. 

The  Chemical  Nature  of  the  Substances  in  Which  Purine  and 
Pyrimidine  Bases  Exist  in  the  Animal  Body.-  In  freneral  it  may  be  said 
that  the  amino  jiui'Imcs  iiilcnine  and  guanine  fos;cther  with  the 
jiyrimidine  bases  tliviniiie  and  cytosin(- occur  combined  with  plios- 
l)horie  acid  and  a  carbohydrate  in  the  vjuious  niirlrir  acidn,  each  of  whicli 
is  ayrain  comliincd  with  some  sinii)lc  i)rotcin  to  f'oiin  nuclcin,  the  essen- 
tial constituent  of  the  clu-omatiu  of  the  nucleus.  One  of  the  oxypurincs, 
iiypoxanthinc,  may  also  exist  condjined  with  i>ho.sphorie  acid  and  carbo- 
hydrate to  form  a  substance  pi'osent  in  ii;uscle  and  known  as  inosinic 
acid.  The  ^'cncral  scheme  of  ci.i '.Ruction  of  .-i  mieleie  acid  of  animal 
orifiin  is  illustrated  in  the  followiufr  l"ormula  su^r^'ested  by  Leveiie  and 
Jacobs:  '■* 


no 


\ 


/ 


\ 

/ 


I'O 


^ll('x'ls(■) 


(Ml. NO 

( ;;ii:OlilU'  J 


.M,|'1 


no 


\ 


O 

I 


\ 

-    1 


I 

I'O-  r. no      r  II  \o 


(lii'Misi'>      I  Ili_\  riiiiii'  i;riiu|ii 


HO 


]|0 


\ 


\ 
I 


ro 


<Vii.o. 


r  ir,\o 

)     ^•vlllsiti(• 


;'■""' I' 


iro 


I'll. 


s|i|i(itii'  :ii-i.l 


O 

\ 

\ 
O     -    I'O 

/ 
/ 

no 


r',ii,o,  -    f'.H.x, 

i  lit'xos(>>      (  ndt'iiino  group) 


Accordiiij?  to  this  formula  nucleic  acid  ma.v  be  considered  as  a  com- 
pound of  polyi)hosphorie  acid,  coiitaininp:  carbohydrate  groups,  which 
serve  to  link  the  i)liosphoric  acid  molecules  to  those  of  purine  or  pyrimi- 
dine. In  nneleic  acids  of  animal  origin,  such  as  the  example  given 
above,  the  carbohydrate  is  a  hexose,  (i.e.,  contains  fi  C-atoms),  -whereas 


i 


il 


I 

"I 


--■_ 


f  (i 


fjil'l'i 

I;  If 

<-'i 

!  .        ■    f 

1?                   ■  ! 

r  1 

<'!l 

1 .  1 

ff 


if 


,  J:   '  :  I 


:  mu   h 


,1.  i' 


it  - 


I.!  ' 


r:   ■11 


Mii     :  i 


hi 


i 


li  l\ 


(i:is 


MDTAIIUI.ISM 


in  tlins..  of  pl.-mts  (0.  ^r,.  vriist^  it  is  a  iviifns<>  If)  C-iitnins).     Tt  lias  l,r, 
f..uii<I  Mcn'ssaiy  f.  iiitiodn.-c  s(.iiu-  fciins  t(.  <l.'sisri,at(.  tlic  .iinVrcnt  |..ii; 
<.f  the  iiiiclcir  aci.l  iiin|,.rulr  :  tliiis.  Hi,-  wlioK-  inolcciil.'  is  rail...!  a   /,//,^ 
uiirhoti,!,.   ,.a,-ii    i.n.nniiiK-lcnti.i,.    ni..l,...ulf    „f   uliiHi    is    ..o„i|,„s,.,l    „r 
I-hosphcri..  ari.l   „i.,|.v,il..  ,,li,,  a   niuUo.i,!, ,   winci,   a?aiii   is  eomp..s,.,l   ■ 
a    imrinc   ,>v   i.yiimi.liii..    miclnis   atta.-li(>(]    to    pontosc   or   Ii.-xoso.      Ti 

nu.'l.'osi.l,.  is  so  nanicl  1 aiisc  it  is  similar  in  stnictiiio  to  a  s;1ii.M.si,|, 

Apart    from    .Iiir..r,.n.'..s    in    tlio   .-arlioliy. Irate    -loiip,    it    ai.p,.ars    tl.;  ■ 
tli.'iT   is  a   rlos,.  similarily   in   tlic  stniclnrcs  of  nnclci.-  acids   from   .|i' 
f.-n-nt  •■.■lis.    Tliis  wonl.l  in. Urate  a  c.mimon  function  for  tli.-m  all    «lii.- 
iriny  W  ..itli.'r  of  a   skeletal   or  of  a    i)liysiolo;ri,.   nature;  that    is,   nuel,., 
aei.l   may   liave   to   ,1,,   nitli    the   snstentaeiilar   material    that    l.n'ihls   il  . 
nueleiis.  or  it    may   have  to  ,lo  with   some  pl.ysiolo^rj,.    fnnefion   eomi,,.,,, 
f"   all   eells.   sneh    as   irritaliil-.- .   or   -rowth.    or    respiration.      If   ,niel,.: 
nei.l   ,s  merely  a  snstentaenlar  material,  then   the  stn.lv  .,f  the  hehaxin 
of  ehnm.os.imes  ami  ehnmiatin,.   in   e.-lls  ,-an   not   hav.-  the  si-nifi.-anc. 
that  It  woiil.l  have  nere  iiiie',.ie  aei.j  .•on.M.rne.l   in  th.-  m-ire  vital  aeti^ 
itics  of  the  ini.-lens.      All   the  mmmH,..]   niieh.ar  stains  .,we  their  s,,,,-!!, 
stainiii-  properties  t,.  the  fa.^t  that  they  ar..  of  a  l.asie  natnro  nii.l  ,•.„:, 

Iiine  with   nnel.'ie  a.i.l.      Cntil   we  know    mor,.  .lelinitelv  what   tl xn,-. 

funelion  of  iii„.l,.i,.  aei.l  may  I,..,  it   is  nnuise  to  |.laee"too  miieli   wi J  • 
on    th.-   hehavi.ir  .,f   the  ehrom..soii„.s   in   eyt..looie   ivsea relies. 

The  History  of  Nucleic  Acid  in  the  Animal  Body.  \V,-  shall  li..' 
-f  all  stii.ly  th."  manner  in  whi.^h  niiel,.ie  aei.l  may  h,-  hn.ken  .l.nvn.  A- 
IS  to  he  expeet.Ml  from  ils  eompl.'x  strii.-tnre.  vari..ns  Ivpes  of  enzMii.  ^ 
•■"•e  .MnHMMne.!  in  this  j.roe.'ss.  The  first  to  aet  are  known  as  the  »n„  ', 
«s,s:  They  split  the  t.'t  raimele..ti.|e  moleei.l,.  into  two  .liiuieleoti-h  . 
uhieh  imm.Ml.ately  aflerwar.l  split    fnrth,.r  int..  monoiiiiel,..,li.i,.s.      l-\„r 

MU.'l..,ti.|..s.  tw,.  of  pnriiH.  an.l  lu,,  .,f  pyrimi.jine.  aiv  tluis  f..rme.l   IV 

eaeh  mo|,.,.nl..  of  niiel..!.-  a.'i.l.     Ka..h   nnel,  .,ti,i..  m.,l..enle  mav  n..w   u: 

•''■'■-"  •' "'I'"^ili"n  in  o. f  tw.,  ways:  <],   ,.itl„.,'  I,v  th,.  spliitin-.  ,. 

of  ph.,sph.nie   a.-i.l.   I..avin.,r  a    nuel...,si.|,.   innanosin..  'or  a.l..|i.,siMe'    ,, 
(■2.  I>y  the  split. in._'oir.,fl,.,ili  pli..sph.,rie  aei.l  an.l  earl...hy.lrate,  lea\V   - 
free  imrine  has.^s.     \,u!n,s,s  hav,'  I,...,,    f.uni.l   whieh   sp..,-i(i,.aiiv   vftV. 

either    .,f   th.vs,.    .{...■.,mp.,sit  ions,    an.l    th.'y    hav    1 „    eall..,l    i.h.,spl, 

nueleases*  (I,,  an.l  imrii,..-nwel,.ases  d'^,   resp,.etively.      In   the  .!. i.ip 

^" '■  "'"■''■■'■  ••"'i''  "11  "I'  'II"  fonr  piir .•..mponn.ls     .,'ii,-,nine,  ^nia„ 

sine,  a.l.Miosihe  an.l  a.l.nin..      may  he  ton 1.     This  is  illiisl  rat.'.'l  In  l! 

"'■'■"""•■•'">'"«   ^••'"■'"■•'.    i"    «lii'-li    th..    nnelei..   a.-i.|    is    r..pr..s..nt...|    .-is 
imriii..  iinel...pti.lt.: 

•Tl,,-  Tn„„l„rs  r.  f,r  I..  Ilir  ,  „/v,„.  .   I„.|i,,„,,i  ;„  ,|„.  .,  1„  n.r 


■'i    ! 


ruic  ACID  AM)  Tin:  iMKiNi:  ii()i>ir.s 


can 


NucleUAeid  (without  the  pjrinidine  group) 

(2)«-'(l)  (T)        ~^(2) 

/  (^Action  of  tiueteosea) 

Ouaninef-(7)  Ouanosiae  Adenosine  (8)— ♦Hdenine 

(3) 


\ 

•*A.ae 


(4)  (5) 

(Action  of  deaminiHng  enzymes) 

Xanthosine  Inosia* 


(6) 


(9)  {Action  of  hydroly:\ng  en:yme»)  (\(^) 
UrieAcMt-(ll)  Xanthine  « (11) ♦  Hypoxanthine 


(Jrtton  of  xanthine  oxidase) 


(Jonei.) 


Til'"  iu'\f  step  ill  tlic  ilisiiiti'tri'iitiiiii   procoss  is  lliiit   tin'  aniiiio  <rri)up 

is  ic.     )\t'il  jihil  til )nt's|)iiii(liiiu'  ii\\  piliilic  is  piddiiccil.     To  brill);  this 

:iliiiiit,  tlicic  exists  :i  spi'cilic  ili  iitiiliii:iiiii  t  iKiiim  lor  cacll  of  the  iihovc 
aiiiiiio  ••oiiipoiiii<ls.  and  each  i'ii/.\iii('  is  iiaiiicil  accordiiiji  to  the  exa<'t 
amino  piiiinc  upon  whicdi  it  acts;  thus,  niianasc  i:!),  ^'uanosiiie-dcaininaso 
(4^,  adciiosinc-dcaininase  I'l'i,  and  ailcnasc  llii  have  all  hfi'ii  id  ••if itii'd. 
The  Irce  liase  may  then  lie  split  olT  Troiii  the  lUK-leosiiles  1  specitie 
hiiilrolif.iiiii  I  ii-.i/iiii  s  {~i  )   (,S)   (!l)   (10'. 

The  .joint  aetion  of  tln-se  enz.vmes  leads  to  the  formation  of  oxypui'ines. 
xanthine  and  livpoxanthine,  which  are  oxidized  to  uric  acid  hy  .rnnthhif- 

ll.tlllllSI      (111. 

In  man  and  the  aiithi'o]>oid  aiies  uric  acid  is  the  end  product  of  tlie 
above  eliaiiires.  hut   in  other  mammals  most   of  the  niic  acid  is  fuither 

oxidi/ed  into  alhintoiiie.     It  has  also  1 n  found,  except  in  man  and  the 

(•him])anzee.  that  extracts  of  oij^aiis  such  as  the  liver,  are  capable  of 
(lecoiiiposiiii,'  uric  acid  into  allantoine.  The  idciititii'ation  of  these  speeilie 
eiizxiiies  is  souirht  liy  a  determination  of  the  free  amino-i)urine  liases 
and  the  ]iliospl|oric  ,ieid  iirodneed  liy  allowiiiu;  an  acpieous  extract  of 
the  tissue  in  (|uestion  to  act  on  nncleii-  ai'id  lof  yeast  i'  at  body  tempera- 
ture. Another  jiortioii  of  the  diL-csteil  mixture  is  then  hydrol\zed  by 
means  of  boiliiiLr  sulphuric  ai'id  and  the  eonstitueiils  ai-'ain  determined, 
from  the  i-esnlts  it  is  often  possible  to  draw  conelusioiis  as  to  the  exact 
nature  of  the  en/ynies  ])resent. 

The  most  remarkable  outcome  of  thi^  work  has  been  to  show  that 
Ihr  ilislrilniliiiu  III'  till  (;i://m(\  /\  mil  llu  siiiin'  in  tin  tlxmirs  niul  orijaiix 
iif  iJIffrrnil  aiiinutls.  \'ery  brietly,  some  of  the  most  imp<irtant  results 
that  ha\e  so  far  been  obtained  are  as  follows:  (lastiic  and  i>aiiereafic 
juices  iln  iii't   contain  a  trace  of  ■,\t\y  >■(  the  eir/\  nies      Intestinal    juici', 


<W^i 


•VtM-t  nmlcic  mill  is  ii'.t'i!  1himii--c  it  i--  Ic-.^  if>i>|.iiit  to  -li-'iiilfyiatiiuj  ih.iii  lliyniir  iinctt' 


'Url 


i  ''  I .'  i 


!        I 


11 


ii' 

i 

.J   I 

-   4i    ■ 

^  i 

i 

•  I  \ 

(i-U) 


MKTAIIOI.ISM 


on  the  othrr  han-I.  ......fains  .1  imeloasp  capable  of  splitting  the   p,.lv 

nu.-hM,t.,l,.s  ,„t„  ,.,„„„.,„..l,.„.i,l,.s.     Th..   twn   py.-i...i.li„o   „.„.l..„ti,I..s  s,,i„ 
"fi   -lo  Mot   ,i...l,.>tr„  fi,,,|,p,  ..iiai.j.'..,  iM.t   th.'   ,„..'in..  ..ii..l,..,ti,l..s  an.  .•„„ 

vert.'.l  lilt,.  iiii..l,.,.si,h..s  (the  .■.izyiii,^  hein;:  .lesi-natcl  "iiu.-h>..ti.h.se" 
Kxt.7i.-t  ..f  the  iiit.'sti.ial  in.i..,.sa.  i.esi.les  haviiif?  the  same  n.-tion  as  11m 
ii.t.'sti.ial  jiii.-e.  ••an  also  tlee.miix.se  the  piiri.„>,  I,„t  not  the  pv.-iini.li". 
Mi..'l.^osi,h>..    i„tt,   ..a.I..,h.v.]fa1e   a.i.l    i.,..'i,>e    srn.ups    (speeilie   a,-li..n   .. 
••'iii.-h.,Ksi,las,.'M.     A   sii.iila.-  a.'ti,..,    is    pi-„.|,„.,.,l   l.v  ,.xt.-a.-ts  of   ki,!..,.\ 
heai-t  niiisele.  a.i.l  liv.M-.     P,|,.o<l  seiui...  h.'i.K.lvze.l  'hl.,o.l,  and  extra.^l  ,, 
pnnereas,  on  the  ..tl.er  ha.i.l.  a.v  eapahle  ..nly  ..f  eanyi..i;  the  .leeo,..p  . 
sition  as  far  as  the  n'.()f;(intii'le<.ti(les. 

Rearnnlinj;  the  other  enz.Mi..'s  mentioned  !..   the  ahove  list,  it   is  i..i 
p..rfa..t  t..  n.,te  that  th,.y  appear  at  .lifTere.it  sta-.s  in  en.I.ry..nie  develop. 
KO'iit,  and  that  Ili..ir  .list.>il.nti..ii  varies  eonsidei-ahlv  in  differe-.t  speei.s 
of  a.lnlt  animal,  the  splee...  liver,  thymus,  and  panercas  eontaini.u'  th...,i 
most  abundantly.     The  disti'ib.ition  of  enzymes   in   the   orirans   of  tr 
m<.nkey  .•esen.bles  that  in  the  b.wer  animals  e....sid.Tablv  more  than  i« 

d..es  that  in  man.    Some  .■emarkablo  faets  have  eome  to  liudit  re-'ardhi- 
puannse  and  adenase.  partieularly  that  suanase  is  .lefieient  i.i  the"orf.nns 
of  the  p.fT.  in  the  urine  ..f  whieh  animal  it  has  also  b..en  f.nind  that  th.^ 
purine  ba.ses  a.e  in  exeess  of  the  ui'ie  aeid.     This  absenee  of  .-uanns.. 
no  doubt  aee..unts  for  the  faet  that  deposits  ..f  (.M.:...ine  n.av  oeeur  in  11. 
n.nseles,  and  that  these  may  be  so  lar-e  as  to  e..nstitute  the  eonditi-m 
know.,  as  guanine  ffout  foun.l  in  this  animal.     Adenase.  o.,  the  oth,- 
han.l.  IS  absent  f.om  the  or-ans  of  the  rat.  Mhieh  a^ain  e.rresponds  with 
the  fa.'t  that,  whe.i  adenine  is  injeeted  sul.eutai.e..uslv   into  these  nni 
mals,  It  undergoes  oxidation  without  the  removal  of  its  amino  ffr..up 
In  the  human  organism,  adenase  appeals  t,.  be  ab.sent  from  all  of  th.' 
organs,  whereas  iruai.ase  is  pr.>s.t.t  i.i  the  ki.lnev.  hu.g  and  liver    bn* 
n..t  in  the  pane.Tas  or  spl.^en.     Xanthine-oxi.lase  exists  o.dv  in  the  liv.r 
The  distribution  of  uriaisr  is  i.eilini,s  the  m..st  i.iterestins     It  is  pres 
e.it  .n  most  of  the  h.wer  animals.     On  aeeount  of  its  presence  extract, 
of  the  hver,  spl,.en,  etc..  i.,  all   breeds  of  .b.jjs,  with  the  exeepti..,,   „• 
Dalmatians,    rapi.lly   destn.y    uri..    aci.l:    a.id    practicallv    no    uric    a.i.! 
Mhen  inject.Ml  subcutaneously  can  be  recovere.]  unehan-ed  in  the  wuv 
but  appears  as  alla..t.,ine.    Tricase,  however,  is  abse.it  in  man     This  h,-,. 
been  demonstrated  by  findinir  fl)   that  when  uric  aeid  is  injected  suL 
cutaneously,  nearly  ail  of  it  appears  in  the  urine,  an.l  (2)  that  uric  a.^iJ 
is  not  destroyed  when   extracts  of  the  organs  are  incubated  at   bo.h 
temperature  with  u.-ic  acid  or  its  precursors.     It  mu.st  of  course  be  k.^t 
in  mind  that,  although  the  uiic  acid  is  thu.s  shown  not  to  be  destrov..! 
in  vitro,  it  may  nevertheless  be  destroyed  in  the  living  animal. 


\m 


uic  Aiii>  ANi>  Tin;  ri  him:  r.(ii>M:i 


041 


Tl. 


il   loiilts  rests  ill   tlip   fnct   lliiit 


he  iiii|Hirt;iii('<'  ol  tin'  mIium'  iltsciiln'il  roiills  rcsis 
fmiii  tliiiii  uf  iiiiiy  liopc  tn  lie  !il>l<'.  ultiiiiiiti'ly,  to  stiitc  csai-tly  in  wliiit 
(Piiriiiis  iilnl  ti>sM('s  tlic  iiitciiMKJiarv  iiict.ilMilic  pioccsM's  (•((iiccrni-d  in 
nucleic  iicitl  inctnlHiiiMii  occur.  Tlic  w.nk  at  tlic  present  time  is  of  sjie- 
cial  siLrnilieaiu'c.  since  it  rcpicsi'iils  unc  type  nf  evidence  wliicli  wi"  iiinst 
liiive  IpcI'chc  we  can  trace  exactly  every  stc])  in  the  iiietalioliMii  i<{  any 
iitlier  liioelieiiiical  sulistaiice. 

Tlie  aliseiieo  nf  uricase  fnnn  liic  lissu t'  man  |)laees  liiiii  in  a  uiiiciiic 

position  witli  reuard  to  tlie  inelaliolisin  oi'  nucleic  acid,  and  renders  the 
investic-ation  of  tlu'  pioliiciii  i)arlicnlarly  difticult,  since  animal  expei-i- 
inentation  is  useless.  Recently,  liouever,  S.  I{,  Ueneilict  lias  discovered 
that  the  Dalmatian  hrecd  of  d<iji  also  known  as  the  carria^rc  dojr,  and 
liavins;  a  s])otted  or  niotlled  skin  has  a  i)uiiin'  iiietaholism  like  that  of 
man.'  Whi'ii  fed  on  food  containiu!.'  no  pniine  suhstanees,  he  excretes 
lai'^e  (|Uaiitilics  of  iirii-  acid,  and  when  the  latter  siilistanee  is  injected 
suliciitaiU'Diisly.  it  is  elimiuati'd  cpiant itat ively  as  sui-h  in  the  urine.  We 
shall  see  later  how  experiments  on  this  animal  have  hecn  niaile  use  of 
in  till'  in\esrn;ation  of  prolilems  of  puriiu'  metalio|i>m  as  apjilicd  to  man. 
In  all  other  animals  most  of  the  uric  ai'id  is  oxidized  to  allantoiiic  liefore 

hciiijr    excreted.      The    dciiice    to    which    this    -iirs    varies    hetweeil    T!> 

and  !•><  per  eeiit  ol'  the  uric  ac'id  in  dil'ferent  species.  This  i.as  heen 
callcil   the   uricolytic    imlcx    illiuiti'r  and   (lixfiisi. 

The  Balance  between  Intake  and  Output  of  Purine  Substances  under 
Various  Physiologic  and  Pathologic  Conditions.  The  main  purine  ex- 
cretory ]nii(luet  in  mail  is  uric  acid,  hut  thi're  is  also  a  certain  amount 
of  purine  liases.  The  |)resence  of  uric  acid  has  atti'acteil  attention  for 
a  i.'1'eat  man\  decades  in  medical  invi'stiiiation.  Iiecaiise  i>{  the  relati\e 
ease  with  which  it  can  appioximatelx  lie  deterinined  ipiantilativel.v,  ami 
lii'cause  of  the  well-known  fact  that  it  m:i.v  In-  I'csponsilile  for  certain 
diseases,  such  as  L'out.  when  it  accumulates  in  tlu  tissues  in  an  iiisolulile 
I'orm.  On  a  diet  containinii'  meat,  or  more  particularly  on  one  eon- 
tainiiiLr  lilamlular  suhstaiices.  the  total  dail\-  excretion  of  urie  aeiil  is 
Very  coiisiderahly  liieater  than  when  the  diet  contains  no  such  food 
stuffs.  The  <'onelnsion  which  iiuriaii  ami  Si-hiii'  ilrew  from  this  oh- 
scr\ation  is  that  |)urine  must  he  jiartly  of  i  .nii/i  hhkx  and  partly  «f 
I  ikIiiiii  iiiiiin  oritrin.  in  othei  wcpfds.  some  of  it  is  derived  more  or  less 
diiM'ctly  from  |ierformed  ptirine  suhslances  in  the  food,  ainl  the  reniain- 
der  fr<im  the  purine  constituents  of  the  animal's  own  tissues. 

Endogenous  Purines.  It  was  thon;.lit  that  a  <lerinile  proportion  of 
e.'ich  of  the  administered  purines  could  he  in\ariaMy  reeo\ered  from 
tin'  urine.  Aithou'4:h  this  has  not  hecn  fi>uiid  lo  he  e\acli\  liue.  there 
is   nevertheless   a    certain    coiistaiu'y    ill    the    jiroiiortion    id'   administered 


.::!]'.  1  [. 


I  i 


V 


'I 


y  f  111 


8 


% 


r   H 


' « 1 1: 


(»    ! 


^'r        ^ 


i: 


it    I 


■I' 
is] 


m 


iV 


III 


(i42 


.MITAIIOI.IS.M 


purine  that  is  oNcvf.-.l.  Thus,  M,.,..!.'!  „n.l  Lvn.an  hnw  fonn.I  rcTntl 
ll.a!  alM.iit  (Id  p.',-  ,.,•111  of  i.i.i«'<-tr(l  li.viK.x;uitl.iii(>,  HO  ,.,.,■  c'lit  of  xai 
Ihiii.-,  1f»-:!n  |„.,.  ..,.nt  of  -uanosii,,..  an.l  :!(l-:!7  ).,-,•  .•.•nt  ..f  a,ln,i,u.  m,,. 
•■Ii'miiimI.mI   ;,s   mi,.   ..in,!.      WIh.i,   coinliiii...!    i)i,i-in.>s    -].,..,   nu.-lcar   ninl.^ 

'■'■■''     ■"■'■  -''•■"•  ""!>■  ••'  '^•"•••'l  !"• x-ti-'M  of  tl.<.  |„„i,u.  rcapprars  as  uip 

.■•'■I'l   111   tlu.   uiMi...      TI,...T   is,   tl,..,vfo,o,   ,.,    .,,„.,.,!    i.arail.-lisn,    hrtu,.,, 

""•  '""" '""•'"  ">'  ""•  ''"'"1  -iiKi  tliat  of  ti,..  nrin...  Ml.iH.  indicaP's  tli.v 

'""""'■'"•''   '■ 1  '^'"'"''1  '"'  -'liininaf,..!  f.on,  tin-  .liH  of  pnticits  wlw>  ai. 

suir.riMir   f,,„„  ,i,.,,ositioM  of  ImsoImMo  iirjito   in   tin.   tissurs,  ,,s   in   ,t 

Tin.  iMl..  of  tiic  p.uii,..  tliat  <lisapp..a.s  i„  tlu-  ImmIv  is  unknowM-  somh-  m 

"  '"•'>  '"'•' "'I>-'^''<I  ill  the  intcsti,,,..  I„il  «l,v  so  ,Mu,.i.  of  il,,.  n.niaiu.lci 

.-itl.M-  al.sorptio„  l.y  III,.  i,loo,l.  slionM  .lis,,,,,,.;,,,  is  a  iuvsPtv.  sin,...  „„ 
iii-i.-as..  ,.,,„  1m.  .1is,-ov,.,v,1  in  ;i,iy  of  II.,.  ,.r-ans  or-  tissues.  Tlie  ,I,.sti„x  ,.,1 
l>uiM„.s  ,.Mn  not  1,..  sl,o^^n  to  intlu..n..,-  any  of  the  other  w,.li-kn.iun 
iiitr,i>;,.nous  nietaliojitfs  ,if  tin'  iirin... 

Th..  foliouini:  tahle  ,.f  ex|)eiini,.nls  l.y   Tayh.r  an,l   Rose*'  n.av  s,.,v,. 

t..  illnstrat,.  tin-s,.  points;    The  snh.i,...t  «as  pia,-,..!  on  a  pnrin,.-f,' |i,, 

-•nnsistnm   of  milk.  e-'s.  slareh   i,n,l  sn^rar.   f,.r  thr. ;,vs       \ft,.,.  thi. 

'"■'•'"■'  ■•'  1'="-'  "f  '!"■  '"tal  nitn.,n.n  ,:!  ,M-a,ns,  was  suppli,.,]  ;,s  s«,.,.t 
'ii-a.ls  th.xnius  Kh,n,|.  el,-.-  eontaininu'  a  hi-li  p,.,-,.,.nta!.'e  (()4S->)  ,,1 
P'l'iiii'  nitroir,.n:  to,-  another  ,.,.rio,|  of  fonr  ,h,ys  still  more  of  the  nitr.. 
;rrn  m;  jrrainsi  was  r,.pla--,.,l  l.y  su  ,-ell.r,.a,l  nitn..,'en-  ai-1  this  avis  fol 
lmv,.,l  l.y  ,.,  |in;il  p..|.io,l  i„  ,vhi,.|,  the  orifri.ial  .liet  of  „,i|k.  ete  „  illi,,,,, 
Pii-iiie  snl,slaii,-,.s.  was  rest,.re,l.     The  f,.ll,.wintr  tal.le  -ives  the  n-snll. 


1st  PKiaoii 

rna\K-Ki:i:K      I'm.  numi,       ;:i;i.  i,:i;,oi,      ,., 

I.IKT 


rrc;i    N    ,-,i„r   S\\ 
<  'iral  iniiic 
I'liriiii-  \  ,  tn(:i|  I 
I'ri.-  :i.i,|  \ 

KrllUlill.li'l-    \ 


4TII  I-Klllol. 

aoi.      I'liMM-M-iai: 

IllKT 


s  't 

_ 

,s,s 

'■" 

7.1 

:.i 

7.i>r, 

11. .■.."> 

II..".,; 

e  (7 

0.11 

e.i: 

e.:.',: 

11.10 

(t.il'l 

II.II 

II -1 

(1 117 

(Is.; 

l.ls 

l.ls 

.''"'"'  ''"•'■'■"^"  "'■  '"■''■  "••'"'  " "'it,.,l   for  |,.ss  than   half  of  the  ..nrin. 

""■•n.^.-n    .n^..s„.,|.     This  ,-,pp,.Mr,-,l   ..s   mi,.  „.i.|.   ,|„.  ,,,,,,,;, ,-         .;,„ 

I'a.s.'s  l.,-in^r  |.i-a,-li,-,illy  nni-haiiir,.,|. 


CIIAI'TKK  I.XXIir 
rUIC  ACID  AM)  TIIK   I'llMNK   I'.ODIKS   (ContMi 

SOURCE  OF  ENDOGENOUS  PURINES 

Kvfii  jift.T  111,.  ,.|itir..  i.Iinii.iiiti.m   .,f  .-ill   puiiiio  siilistjii s   hum   1li(> 

f<>"<l  ill  flic  i-ASi-  uf  iiiiiii,  i)iirin.'  (•(.iitiiiucs  t„  1„.  ..xcrctcl  in  tlio  iiiiiie 
!is  uric  iici.l,  Tliis.  ;is  !ili..\<"  tviiijirkc.l,  is  called  cii<I..u'cii..iis  excretion. 
At  first  it  \v;is  thoiiulii  l,y  H„ri;,ii  iin.l  S.-lmr  lliiif  flic  tot;,!  iiifro.r,.,,  of 
tlic  iM.rinc-fVcc  .lict  ,.,„.M  I,c  co,isi.]cr,-,My  v„r.c,l  without  cniisiMu'  iiiiv 
Mltcrafioii  in  flic  iinioiint  of  tlic  ..ii.loyciioiis  luiriitc  excretion,  l.iit  a  rc))- 
etition  of  the  work  has  slwnvn  tli.-if.  wliei,  these  ch.iii-es  arc  of  coMsi.ler- 
ahlc  niaf,iiitii.le.  the  ••n.losr,.,„.iis  ni,,i,.ty  ,l.,cs  not  remain  constant.  This 
lias  alrca.ly  luvn  .lenionstrafc.l  in  the  tahlc  on  Folin's  results  fsec  pa.rc 
<il4).  an.l  is  still  l.ctter  illiistratd  in  the  a.-coMipanvinjr  table,  which 
.shows  the  excretion  of  uric  aci.l  an.l  eoi„ci<lentIy  of  urea  from  liour  to 
hour  in  the  urine  after  takiii!.'  food  which  is  free  from  nudcin  or  imrino 
suhstances.  After  a  fast  of  six  hours,  a  diet  consisfintr  of  hrcad  and 
potato,.s  was  faU..n  at  !::!().  and  the  urea  and  uric  acid  measured  in  the 
urine  eacli  hour  thereafter.* 


■n.iiK 

ri!K.\ 

I'KIC  .Mill 

.\.MOI 

.NTlIf 

riti.NK 

CM. 

M<;. 

e.c. 

10  11 

i.e: 

•2>'> 

11  IL- 

1.1.-! 

27 

IIS 

1-1    CM. 

i.iir 

LM 

MU 

1  J    (  nn':tl) 

o.i;  t 

21 

<;o 

1.1-' 

.>.» 

4:! 

111! 

.'IS 

41 

t  .» 

O.St 

40 

5.1 

.')  (i 

I.Hi 

."i<! 

nn 

1.20 

:'.!> 

5(i 

1..17 

.•!0 

n.j 

1.»7 

;'„■■ 

1S.1 

Hid 

mil 

1 .:!:; 
l.:r, 

L't 

1.-..T 
ISO 

1  II"iikiiis  mill    liii|i('.  H'i 

^1  post pra  11(1  ial  iiirnm^f  of  (iiilonnions  imrinr  (.rcnlioii  is  verv  dis- 
tinct, and  if  indicates  that  rluriiii;  the  priK-css  ,if  as.siiiiilatioii  soiiiethintr 
must  l.c  ,„.,-urrin^'  in  the  <*rf:a!iisiii  \\hicli  entails  flu-  pr.Kluciii.n  of  juirinc 

..f  :j:^i  T:;:i:;??:r;K,^l;;tL;;::,  r'';r:;::;.;ri;''::'i.,;;,:;;;-  t:t.u:  •ii.r::::;::,:';;; •-'"' 

043 


IFi 


i   .  i! 


i  ■ 


ll 


r  '1 


;  r 


fi44 


MKTAIIdl.ISM 


''"'"  ' 'tfiiism  ils,.|f.     As  to  ,vl,at    lliis  ,„;i>    I,,.,   if   is  impossil.lr  |.. 

say.      It   iiiny  1...  iissn.-i,,!,..!  ,vilh   ti,..  woik   of  tlic   u'iistri,.  i,ii.|   iiitrsl  in;,! 
Kitlll.Is.    uliirl,     m-dlls     li„.     iiitcvsliliir     MljiK.stiHM,     ..Iij;i„;,|lv     ,n,„l..     I, 

ll..rl.M.-/..«ski,    ih.it    in,r,,s,,.,|    sul.st.,u...s    i,„.,v.s..    tlw    ..N.-nVinn    ,.f    „,„ 
n.-Ml  l.y  ..,„isMi..'  a  l..i.,-.M.y|.,sis.  tl,..  ,,nri.,..  iMin-  .Irriv.-l  iVnin  ||„.  ,Mi,.|,.i,. 

'■"■'"'     ''"■'     '■"■''''     "'"'"     ""•     ''•" Vt.s     l„....,.nC     l.lolx...,     .I..VM1.        TIlMf     tllis     i. 

""'   •'" '■'■'■'■'   <'NpIanali..i,.   h.-wrv.-r.   is   in.li.-Mtr.I  l,v   tji.-  ia.-f   lliat   in 

!.'.'sl,..l  sMlKta,i,-..s  tl,„f   t:iv(.  i-is..  t..  an   iiwiriisr,!   imnilH.r  „r  1,., vt.s 

alT,.,-t   tl„.  i.x.T.tioM   ..r  ,iii.-   .M.-i-l   ,h,r;n„   ft,..   ,,,.ri.Ml   ft,,.   l,.i.r,M.vt,.s '  .•iiv 

'"■'''•■'"  '"  ""■  '•' '•  "•"'  ""t  "/'"•  tiM.y  i.av,.  .iisi,|.|H.an..l.  wiii,.),  wumM 

':;"'■  '"  '"■  ' ••'"''  "'■'•'•  ""•  "'■'"•  ••'•■i'l  a  pnMJuct  .,f  iM.riiir  sui.stan.vs 

lil'<T,Mt,.,l  i,y  tJH.ir  l.rraii.loun.     Tliis  unul.l  in.li.-at..  ti.at  tin-  |Mi,in,.  m,1. 
sta.i.T  IS  a  M.ftalH.ii,.  p,„,lu..|   ,,f  ij,,.  livi,,-  Lmk-.k-n fs  aiul   ii,.t  a   l-ivnk 
'1-'"  .1  pr.Mlurt  ..f  tlios..  tl.i.f  a.r  ,|..a.l.     It  s)„.ul,i  !„.  „.,tnl  ti.at  tl,..  in.Mvas,. 
in  tlir  imstpraii.lial  iirir  ari.l  ..xnvti.m  ,m-,mms  ..arli,.,'  tlian  tiiat   ..f  uir,-, 

'''•'"'    """'•    I"''^-iii^'    '|ii<'''li"M    i<.,-,Ms    llu-    orlnh,    nf   the    n„h,.,,„„u. 

l>i>n>H..  I  ,-,<•  :„■„]  is  til.,  pu.il,..  will,  Avl,i,.j,  X,,.  aiv  must  n,„....,-„..,I  in 
''"•  '■.■'-'■  nf  „,.„,,  ,ii,.i  ,.|„.,„iMry  si,..«s  MS  ti.al  it  i„av  1„.  p,„.|,„.,..l  ,.i||„,. 
I,y  tlic  cxi.lati.m  ,<{  tl„.  I„vv..r  jMirincs-  i,a.i...!v.  of  t],os,.  wi,i,.l,  niv  tl,.. 
n,i,st,ti,..|,t  parts  of  III,.  „„.-l..i.-.a..i,l  niol,.,.,,]..  .,  ■  l,v  a  sv„fi,..sis  of  tu.. 
y-'.'i    .„ol(.r,ii,.s   will,   a   <.a,i,on    .•.•si,!,,,.   .•o„taiMii,-   ti,|.,.,.   .-aiLon    at..,,,. 

'""''■'■  "'■ '"•'-■'li'CMtiy  tw.,  so,i,<...s   tVoiM   u],i.-|i  tl,..  ,.,„lo._n.n.,.is  i.inin. 

.•N.T..t,o„  i„  ,„a„  .nay  1,..  ,i..,iv,..|   :  (j,   s.M,tl„.sis  of  two  tirra  m,o1....„1.  . 
a,,. I  (L'     o\i.lati.,ii  .)f  tl,(.  lowci.  piiri,,(s. 

W..  Mill  ....iivi.!..,.  first  the  possibility  of  synthesis.  I,,  l,i,-.i.  ai,.l 
ivplil.'s  l..-a..ti.-ally  all  tl„.  nitroir..,,  is  ,.x..r.l...l  in  tl„.  )'.„■,„  of  ,„.i,.  a.M 
and  it  IS  ..asy  to  show  that  this  I,as  l.,.,.„  pro.l,,...,!  ii,  tl„.  or-anis.u  ^^ 
h..  synthesis  of  luva  with  cai-l.on-ii.-h  irsi.hn.s.  o<.<.uiii„ir  mainly  in  tl. 
Iiv(.,-.  Minkowski  fonn.l  that  I.y  i-..n,.,vi„L'  th..  liver  from  -.-..s."..  Mhi.'l 
IS  ii  .•on,pan,tiv,.ly  simi.l..  op..i.ati.„i  .„,  a,-..o,M,t  of  an  anast.,i,i..ti,.  v,.in 
h.'tw(...n  tl,..  i..„lal  a,,.!  .1,,.  r..„al  xvins.  th..  mi,.  a..i.!  in  tl,..  nrino  l.,...a„,. 
v.M.y   mark.'.lly  .l....r..as..,|   an.]   ammonium    hn-tato   t....k    its   pla<..>   (pa.;. 

•il."^!.      Si,,..,,  w..  kn.,w   that   ami, i,i„,    i,,   ll„.  animal    ho.lv    is  ..r.linaiil' 

<.onv..,t...I  into  nr..a.  wo  may  .■..n.-l,,.!,.  f,...,,,  this  ..l,s.^i.vath.n  that   s ' 

"""-  '"'^  ' "•■'•'■'•  •"  I'i«.v..„t  th..  syntl,..sis  ..f  uira  into  nri.-  a<.i.l       |. 

.•onfirmation  .,f  this  .•.,n..l,isioii  if  was  .sul,s..,pi,.ntlv  f.,„n.l  that  if  a,,, 
m,.n„im  la.-tat..  was  a.l.l,..1  to  th..  hi.,.,,!  i.,.rf,.s...l  throiiirh  tl,..  'is  lat..! 
Iiv..r  of  th..  sj.M.s,.,  ,„i..  a.-i.l  was  ,,  ,,,lu.-...l  in  th.-  p..rfusi.,n  thii.l  •  F,ir 
tl,..rmo,-..,  v,h..n  l.inls  a„.l  r..i.til.'s  a,-..  f...l  with  ammonium  salts  „- 
Avith^W  .l..f;ia.lation  pr.,.l,i..ts  of  prot..in.  th..,..  is  an  iiu-ivas..  in  tl x 

of  »at..r  ui.h  ,l„ir  f.,c„l  ns  ,!,hVr  nninlaN  '    "'   "''"   ''"   ""'    "^•-   '""I'    '••"■«•'   ■,"a..l,... 


U\{ 


JLi'.!." 


rnif  A(  i[)  ANi>  Tin:  imiiini: 


rM 


•Tctiiiii  i>f  mil'  iirl.!  iiistciKl  uf  uicii.  Kvcn  tliiiiir  wliidi  in  a  inaiiminl 
((•nils  to  i-aiisc  MM  iiicicasc  in  moa  cxiMcliiin  causes  in  liinls  and  icjifilos 
a  similar  increase  in  tlie  excretion  of  nric  acid. 

In  tiie  <■  ily  (lavs  of  research  in  llie  nricai'id  prolilein,  not  inconsid- 
eralilc  mis  .i.cs  were  made  on  ; oniit  of  failure  to  recognize  the  essen- 
tial  dilTerenee   in   tlie   iiu'taliolism   of  urii-  acid    in   liirds  and   niaminals, 


am 


1    the    teiidencv    I' 


'or   some    time   after   the   exact    state   of   all'aii 


diseo\ereil  was  to  cousiih'r  tliat  in  mammals  inne  of  this  synthetic  i>r 


Th 


atter   view,    liowcver 


IS   siireiv    ii irecf,    for   a   n  r- 


ess  occurs, 

l,ini  tniiiiinil  mil  mihi  uf  iirir  ai  lil  ilsilf  },,il  „(  llii  Imn  r  i,iiiilli  hoiliis 
till)  hi  itriiihii-iiJ  hii  stnitliit<i!<  hi  tin  iiiinii iiiiillini  hmhl.  Thus.  Aseoli  aTid 
Fzar"  discovered  that  uric  acid  could  he  made  either  to  <Iisappear  or 
to  he  formed  w  lien  a  miiieed  preparation  of  liver  was  iiicuhated.  depend- 
ing' upon  wJi"tlier  oNvircn  or  carhoii  dioxide  was  huhhled  throii!.di  it. 
With  oxyiren  uric  acid  disap))eared.  whereas  with  earhoii  dioxiile  uric 
acid  aiM-iimiiliiicd.  indicating'  that  in  the  presence  of  this  iriis  the  destroye.I 
uric  acid  hecaiue  reformed  from  tlie  disiiitetiration  products  of  the  oxy- 
t'cnation  jirocess.  As  similar  results  were  ohtaiiicil  from  the  livers  of 
hirds,  it  is  clear  that  no  essential  dilTerenee  exists  hetweeu  the  purine 
metaholic  processes  <iccurriiiir  ii:  the  livers  of  hirds  and  of  mammals. 
Tho  ditTerenc(>  is  a  <|uaiititat ive  not  a  «|"'i''tiitivo  one. 

Resrardint'  the  ciieniical  nature  of  the  product  into  which  uric  acid  is 
hroUcii  down  and  from  which  it  may  he  resynthesized,  it  has  heen  jtos- 
sihle  so  far  to  ich'iitify  hut  on(>  siihstaiice-  na'iiely.  dialuric  acid.  This 
is  a  1  rplexini;  result,  for  from  all  other  iii\  esliirations  it  would  appear 
that  in  mammals,  with  the  exception  ..f  man  and  the  anthropoid  ajies. 
uricase  splits  iirii-  acid  into  allaiitoiiie  (see  pai.'"'  ti4<H.  wiiidi  siihstaiice, 
however,  when  adde<I  to  liver  extract  did  not  cause  any  uric  acid  to  he 
formed;  nor  did  any  of  the  otlicr  know  n  decomposition  iirodiu-ts  of  urii' 
acid  liave  such  a  result.  The  chemical  reaction  involved  in  the  ])roduc- 
lioii  of  uric  ai'id   from  dialuric  acid  and  uira   is  indicated  as   follows: 


XI  t 


()  =  (• 


\ 


/ 


/ 


-r=:0 

I 


II.OII 


II       -Ml 


\ 


Ml  -   (■   -^O 

( <ll:ilin  i'-  :i('iil  i 


II      Ml 


\ 
(' 


The  synthesis  ,,f  uric  :irid  is  hrouudit  ah.,ul  l.y  the  comhiiied  action 
of  a  thermolahile  eii/ym<'  in  the  hlood  and  a  thermostahle  hody  in  the 
ti.ssues.     An  a(iiieous  extract  of  l.lood-free  liver  of  the  doi;  can' destroy 


f. 


I' 


kill 


hi' n^ 


n 


ui'l 


Hi:  I 


!: 

M 

■  i 

•■\ 

f^r^aj 


(ktii 


MITAIIill.ISM 


iin«'  fH'id  mmI.v   in  tlu'  pirs,.!,,..-  ..f  ..n.vu,.|i:  it  ,.aii  nnt   wfuvm  it   rviMi  in 

•'"'  •"''■'^'•' '■  '•"'■' lioxi.l.-.     On    th..  ..th.T   luiii.l.   I>l 1  scntiM   .•iin 

not   ivfoiiM   iiiir  jiri.l,   wli.Mciis  a   ii.ixliMc  „(  tlir  M ll.-ss  liver  .•xtiart 

"'"'    ■'' '   "*'•'""'   I""'li '^   '"•'.•   n.-i.l    -cHlily    iiii.Irr  suit;il.l ..lilioiis 

15..ilinii  of  III,.  liv.T  .•xtniH  .l..rs  nnt  .-ifr.M.i  th-  ivsnil,  Init  l.njlinj;  nf  ti,- 

'•' '    '*'■'""'    '■'•'"1<'''^    it    illrM|.i,l.l..    nr   ..xn'tinK    its   jnint    llrtinn    witii    fl„. 

Iilnnilli'ss  lixcr  r\tr,i('t. 

Tin's."  cxixiiinrnls  witli  .In-'s  liver  servo  only  as  eireiunslantial  evi 
•lein-e  that  nrie  aei.l  synthesis  ,, reins  in  iM.-inini:i.ls  iis  \vell  as  in  l.ir.js 
Afore  ,//,-,, 7  i,n>„f  tinil  i,i,ri,„  siinlh.sls  or,  iirs  In  wniniinls  is  as  follows; 
in    It   XNas  .iiseov.'re.l    Ion-  a-o  Ly    .Mieseli,.r  that   salmon,  fiffer  leMvin- 

th..  sea  I.,  aseen.I  the  rivers,  hj.ve  a   u  ell-ilevelope.!  s.-nlar  system,  hnt 

th.Mt  in  the  npiier  reaehes  of  the  stremn  the  m.isenhir  system  l.eeome.^ 
eonshleralily  alro|.hie,|  an. I  the  t.^stes  en.>rm..nsly  .I.-velo|v.l.  .\s  Ih.. 
lish  t.'ilx.'s  no  f.,.  ,1  .Inrin-  th..  mi-ration,  th.'r.-  mnst  I.e  .•onv..rsi.,n  of 
th..  i.n.t.'in  ..f  III,.  mns,-l..s  int..  the  .-..Ihihir  t>sn,.  ..f  the  sexn.-il  -.'lan.ls, 
••111. I  nm.l.'ie  a.'i.l  i.nisi  1...  |,r.„lne...I.  c.M  .\  h.^n's  e._n_r  |„.f,„.,.  its  ineuhji 
li'Mi  e.-ntiiins  i,ra..ti..ally  m.  nnel.'ie  aei.I.  uh..r..as  after  .1ev..l..i)nn'nt  has 
"ell  start, .1   nM..l..ie  .-lei-l  in<.r..as,.s  l.y  I,.a|.s  :iii<1   l..iun,ls.     Similarly  th,- 

.■u-irs  ,,f  iiise.-ts  inerens,-  in  pnrin t..nt  v,.ry  mark.'.lly  as  .levelopment 

I.r......,.,Is.     c.W    y\]]U   ....ntains   praeth^ally   n.>   purine  .lerivativc.  nn.1  y.l 

uh..|i  it  is  f,.,|  (o  y,,nnir  jrr.'u  in-  animals,  the  or-ans  lav  .>n  purine  siih 

stan.-.s  ahun.iantly.     Tn  .,'en..ral.  in.h',..!.  it  may  h.-  sai.l  that  th m1.in..I 

purine  iin-ivas..  is  in   prop., rt ion   to  th.-  iner..a-..  in  l.o.ly  uei-ht  on   th.' 
milk   .liet.      (4.    In   <  (shorn,,  an,]    .M..n.l.rs  ..\p..rim..iits  aln-a.ly  allu.l...| 
t...   it    has  l,....M   shown   that    a.le.piat..    (..-..uth   .{.p.-iuls   primarily   ..n    tli.. 
iiatiir..  .,f  th..  i.r.it.'in  I.nil.lin-.'  st.m.'s.  an.l  n..t   ii|).>ii  the  imrine  e..nt.nt 
..f  th..   fo.,.1.     (.-.I   An  .,l..i....ti.,n   mi-ht  he   rais..<l  t.,  th.'se   r.-snlts  .,ii   th,- 
s.-or..  that    th..y  .1..   n.,t   a|)ply    t..   thi.   a.liilt    mammal.      lnv<'sti.,'ati..n   of 
th..  |.r..I,le„i  has  hith..rt..  h,.en  s..ri..nsly  im),...1e,|  l,y  th..  fael  that  m.  or 
.linary  lah..rat..ry  animals  v  ,.r,.  known  in  uliieli  urie  aei.j  is  .-xeret...!  ii 
th.'   urine.      The  .lise.iv.ry   that    this  oeeiirs   in    the   Dalmatian   <ln._r  has. 
tH,w..v..r.   lua.h.    it    p.,ssil,l..   f.,r   S.    |{.    |!..„..:|i..t "    t..   sh..w,    not   .mlv    fl,,,i 
after  iner.asin-  the  a.iioiint  ,.f  n,.iipiirine  f..,,,!  th..re  was  a  verv  .listiii.'t 
iiier.'as..  in   the  nrie-aei.1  .xen'ti.,!!,  hnt   also  that   wh..n   the  animal   was 
Uyt  for  a  .vear  .m  siieh  f.M„l ;  th.r.'  was  ..xer..t...l  a  t..tal  am.mnt  of  nri.. 
a.-i.I   at    least    t.n    tlm.s   ureal. 'r  than    .-..iil.!   have   eoiiie    from    the   tra.-.s 
una\.,i.lahly   in.-lml.<l   in   tlie   f..o,l. 

K.-anlin-  the  .•Ii..mieal  nature  of  1),,.  siihstan.'e  from  whi.-li  tlie  pnriii.. 
is  synth.'siz,.,],  w..  know  at  pr.'s.-r.t  jiia.-lieally  iiotliin-  X,,  ,1o„t,|  s,.m,. 
of  the  protein  hnil.lin.^'  .stones  fiinetioiiate  in  this  ..apaeit.v,  pyrimi.liiie 
l).."in<;  i)rol.al,ly  the  pro.lnet  that  is  fi-st  f.)ntio(l.     Tlius.  pyrimi.lino  mav 


IIIC    AMI)    AMI   Tin;    I'l  ItlNI:    iinniis 


f.l7 


lit-  ih'ihIih'ciI  IIS  ii  iTMilt  III'  till'  i-<iinliiii;ili<in  of  iiiniiin  liiiiliuiii-  tiriil  with 
urcii.  fill'  iiiiiiiiiiiimliiMii'  iii'iil  lii'iiii:  prciiliirrd  liy  ruiiili'MMitiiin  of  lij.ilrn- 
•■yiiiiit'iii'iil  iiioli'i'iilrs: 

:;||l'N— .|l\     rilMNi       (di  Ml  1    — .  Nir     fi) 

I  I 

rn  f'N'ir, 

I  n 

Nir  cNir, 

Cliyi1rih'V:iTiir       ('iiiiilM"  iiiiil.iiih'       fiiii'iil      (i'\.v  .liiiniii"  I'l  I  iin'cllni') 
nriil)  iiitiilo'i 

Aiiiilliir  piissililc  siiiirci'  (if  |i\  I  imiiliiii'  i>  tlic  (ixiilMtimi  of  iifiriiiiiii'  t'l 
!:i|jinii'iiir-|iici|iiiiiiii'  iiriil.  wliii'li  lli''ii  rcniili'iiM's  t(i  I'lTiii  Jiiiiiiin  |i.\  rinii- 
iliiio. 

I'liriiii'  sviitlirsis  iiiKliPiilitcilly  ncciirs  in  the  iiiiiiinniiliiiii  Imily,  Init  it 
is  ililliciilt  tci  ri'i'iiiiiiizi'  in  nii'tiilMplisin  iii\  rstiirntinns  liirniisc  it  is  ti  slow. 
riintiiMHitis  pi'iii'i'ss.     Thf  iniiliiiliility  of  its  lurrncc.  Iniw  i'\  i-r,  is  imli- 

I'jltt'd    ii\     slli'li    I'l'snlts    lis   tlicisi'    ilisi'I'ilicil    nil  .  |i,IL'i'    liH.    ill    wllirll    ilHTi'.Isi' 

in  iinrinc  fXiTrticui  is  (iIisim-mmI  jit'tir  v.-iryiii!.'  tlir  iiitii!<t'  of  fi'ml.  I'vcn 
wlirn  tliis  is  itsi'ir  i-ntii'i'ly  d'ci'  ffdni  umiin'  siilistiinci's.     Win'tlicr  m-  ikiI 

rlijiiiiri's  ill  till'  iH'ti\it.\   III"  |iiniiii'  syiitlicsis  iir  in  cciinliticins  df  (lis«'iis(> 

is  ii  <|iii'sti(iii  wliii'li  MWiiits  invi'stijriitiiin. 

The  Influence  of  Various  Physiologic  Conditions,  of  Drugs,  and  of 
Disease  on  the  Endogenous  Uric-acid  Excretion.  Mn^i  ntnr  imri.i)-  wiis 
lliiMij;lit  li.\  r.iiri.'in  to  riiiisc  .'in  ini'ii'.iscil  fxcri'tiiin  <it'  iirii'  aciil,  frnin 
wliirli  lie  ilii'w  tin'  I'Diirliisiiin  tlnit  tlii'  li>  ))ii\jintliiiii'  jircsciit  in  I'nmiwni- 
tivcly  liii'SJi'  iiiMiiiint   in  imisiMilar  cNti-ai't.  nr  its  |iii'cnrsiii'.  innsinic  aciil, 

imist  III'  an  ini|)iirtant  soui'i I'  riiciintcnous  niii-  acid.     OtliiT  nliscrvri's 

(■Ficatlii's,  I'tf.")  liavr  finiiid  tliat  stn  innnis  I'M'i'risc  canscs  a  distim-t  in- 
iTi'asi'  in  iirii'ai'id  I'xri'i'tinn,  \\liirli.  linw  i'\  rr,  is  nnirli  less  iiiavkcd  mi 
ri'pi'titiiin  nf  tlic  sanii'  kind  nf  i'mti-Isi'  nn  tlii'  ni'st  day.  If  snini'  iii'W 
kind  <if  nmscnlar  \Mirk  is  |ii'ifi)inu'il.  aiintiu'r  incrrasi'  in  iirir  acid  will 
vi'siilt.  Tlii'ii'  arc  still  ntlicr  invcstiiratnrs  wlm  deny  tliat  iiiiiscnlar  wnrk 
lias  any  inf  i.'iicc  on  iiric-acid  excretion. 

It  lias  liccn  iiliserved  liy  several  iiivestijrators  tliat  tlie  i'ndo<;eiiiiiis 
purine  cxcfctioii  is  distinctly  hliilur  ihiriiifi  Ihr  wnhinn  Imiirx  Ihiiii  iliir'nui 
shf/i.  This  can  not  lie  shown  to  drpeiid  on  variations  in  the  urinary 
fiinctinii,  and  since  it  is  decidedly  doiilitfiil  whether  ordinary  iniisciilar 
acti\ity  has  any  inllneiice,  the  dinrnal  variation  is  must  difficult  1o 
aceiiunt  for.  The  eiidnuenoiis  excretion  in  man  is  not  the  same  for 
(liffcn  III  lntlirl<hiiil.'<,  even  when  calculated  for  the  same  liody  wei!.'lit  ;  it 
varies  lictwecii  0.12  and  (t.^t)  |ter  cent  iiurino  uitrosreu  in  an  adult  man. 
It  rpinains  remarkahly  constant  for  a  s-'iveii  individual  from  time  to 
time,  lieinjr  unalTeeted  hy  moderate  di'sri'ces  of  viuiation   in  the  amount 


Hi 


! 


II  i| 


1^' 


liH*  I 


w 

t 

j-1 

t;»s 


Ml  T.\IUpr.l>M 


"''  '' '  '••''""■"  l'f"\i-l<<l  lliis  1.0  i.iiiiiio  fm.;  «|i,.M,  liou.'v.r.  til.'  iinmiihi. 

.iiv  I'Nliviinlv    vari^lilo,  .•Ii:iiii.'«'n  ;iiv  |iniilii.f.l   i  «...  |.;nf,.  (l|  Ji. 

Ill   ilis.MM'.    f,  r,  r  . •MUM'S  ill)    ili.'t'.'iiM'il   .'\,.|-.'li(iii,      TliU   liiis    I M    in.isf 

•■l.jirly  slmuii  liy  l.i;illi.w,  «  jm  tnnk  n  hiiu't'  t'liiMiuli  .li.sf  i,(  ;iMtityiili..i.| 

M'liiiii  1m  i.nMJ ;i  ,|i,iii„.|  .li'vfiTc  i.i'  f,.v.'r  i  I(i:i     r.l,  „m<|   f.imi.l  i||.,i 

nil  iiM'icis..  in  mi,' , 'lei, I  rNcivii,,,!  iiin'.I.     Tli.it   iiicnjis,',!  .•m|„IiiisI  i.m 

|>n"-,'».'N   ...•.•iiiriiii;   ill    III,,  tissues   uciv    r.s|i.,iisil.|,.    f,,f   tin-   iiri.'   ii.-i.j. 
Mils  sliMuii   In    til.'  s.'iiii.'  iiiitli.ir,   wild  .•jiiis.'il   ii   siiiiiliir  iih'iciis,.  |,y   siili 
j.'.-tlll!.'  IlilMs.'if  I,,  i;,\,]   |„itlis   (<„■  il   .■.,lisi.i,.|;iM,.   |..i-         ,,r  til,,,.,      'I'll.'    ill 
.'l'.'i|s,',|  loss  ,,{■  li,.;it    thus  jli.l I   sliiilllljit.'.i   tl I.,..     ,lsti.>ll    pni.'i'ss.s    ill 

til.'  Iin.ly  s..  lis  ;.>  iii.'iiiitiiiii  til.'  Iincly  t.in|..r;ituiv,  jiiul  j,s  a  irsult  tli.i, 

«1IS    illl    illc'IV.Is,'    in    mi.'  Il.'i.j    .N.-l.tinll.       It     llllS    Inlli;    Ih'.u    kllnWII    lll.lt    .•ill 

.'x.'.'ssiv,.  iitnniiiil  <.r  nri.'  ii.'i.l   is  ,.\cn't,.,|   in   I,  ,in>,  <,lln  ntin.     Tli.'  iiii.-l.in 

.'I"  i|isiiit.''_'i;it.'.|  l.'U.-...'.\  t.'S  is  .•niiitliuliiv    lirl.l   icspiMisiM.'  U>V  tli.'  ill.TCiis.', 

X.iturjilly.  iiiiii-li   Willi;  Iiiis  li..,'ii  ,i..n d,,,  ,.,i,|,i._ri.| s  iiii.l  cxii'/i'iihiis 

l""'i' N'Ti'tii.ii  ill  '/'.,//.     \,,  viiy  stiikin.,'  iiiiiiiii.'ilj.'s  III'  .'M'lcti.in  lijiM'. 

'"'^* '■''••■•    ' "    l"-'liullt    til    liu'llt.    .MTIll    |i,.|li;,,,s    tllJit    Jlft.'l'   tllC    il,..','stiii|l 

.'f  l>ii.iii.'  rii'li  f,i..ils|iiir>  it  tiiki's  l,iii-,.f  inr  the  nsult'iis;  .'m.k.'Ih.us  .x 

•■r.'timi    til  il.'M'liip  ;i|ii|    |v;iss  ,'l\'  .iv. 

('.it.'iin  <lnt.,s  nn'.'.'t  tl;.'         i.timi  uf  uiii-  ii.'i.l.     Si;li.'yli.'  ii.'i.l  is  s.ii.l 
I"  -•iiiivi'  !iii   iii,.r,.,is.'i|  .'Mifiiiin.  Hill]  I'itr.'it.s  n'l  tiiiiily   Imvc   tliis  .■(Ti'it. 
Ill   lii.tli   .'iis.'s  till'   in.T.'jis.'   is   fiiil.iw.'.I   liy   a   .•..iiip.'iisiitiir.v    fall.  \\liii''i 
iii.iii'at.'s  that  tli.'s.'  .Ini'.'s  a.'t  liy  fiii'i'itatiii-r  tli.>  .'x.-iTtimi  ratli.'i-  tliaii 
I'v  iiillii.'ii.'iiiir  till'  iin'tal...li.'  pi.ici'ss.'s  thai   i..'i    i,  •  s.Min'.«  ,.r  .|,,<  mi.' 
.•ii-mI.     The    'fl'.'.'t  .if  .'jifri'iiii'  has  lien  v.'iy  .'aiffiilly  invpstiirat.'.l.     (iivi  ii 
til  th.'   Daliiiatiaii  .|i.-,',  r..f.'ir,.,|  tn  alu.v.'.  S.   |{.   j";.'!!.'.!!.'!    fnun.]  that   ,i 
small  .li.sr  .-aiis.'.l  a  sli.jht  .l.'.T.'as,'.  liiit  that  a  lai'wi'  .los.'  ha. I  pra.'ti.-all.* 
11"  .'fl'.'.'t.  ;iltlliil|./h   thi'l'i'   \\;is  ;i   m.talil.'   ivt.'iiti.m   .if  liilrntr.'ii.      (>,|    mail 
liiiu.'V.'i-,  .litTiT.nt   iisiijts  wi'i-,'  s.'.'ur.'il,   f.ir  if   was  f..iiii.|  that    wh.n    I 
!.'ram  .if  .■alTi'lii.'  u.is  jrivi'ii  .laily  for  si'vcral  .lays,  a  sliijhl   l.ut  .h'liiiil' 
I>i'.ij;irssiv.'  in.T.'as.'  in  the  ."imIiiu'.'Iumis  iiiic -a.'i.l  .  x.'ivti.in  ..<'.-miv.I,  ami 
it  last.'il  fur  10  .lays  aft.T  the  .'atVi'iii.'  ailministratimi  was  .lisconiinu,.,! 

LilM'ral  all.iwai ..f  this  alkalni-I  may.  thi'ivfuri'.  m,t  |„.  ,p,ii,.  ^„  i,|„o,.|i 

•  'lis  as  it  is  assiiiii.'.l  til  lio. 

Uric  Acid  of  Blood.  In  all  ..f  th.'  invstiLMtiuns  .■i.iisi.l..r,.,I  almM'. 
thn  liohavi.ir  of  iiri.'  a.'i.l  is  .ju.|n,.,l  f,,,,,.  th.'  amount  of  it  .'x.-rotcl  in 
th.'  iiriiii'.  Valiial.l.'  IIioukIi  sii.'Ii  results  must  1.,..  their  iiit.'ri.ri'tati.ui  i- 
always  .lilli.'ult.  sin.',,  two  fai't.ips  tha',  ar.'  <|uito  ii„I..i.,.n.l..nt  nf  .arii 
..thcr  have  t.i  he  k.'pl  in  miii.l  nam.'ly.  the  pioilui-tion  ..f  tlio  iirii-  ai-iil 
in  th..  .,1'iraiis  ami  tissiii's  a,,.|  its  ,'v,.,,.ti„„  !,,..  .i,.,  1,1,1,,,,,.^.  hx  f..t,n.vlinn 
with  th.'  latter  faeti.r.  we  iiiiisf  also  e.iiisi.l.'r  th.'  m.'th.i.l  .,f  transp.nta 
"""  "'■  '"■'•■  "'•'•'  ''.^-  ""'  '•' I  i'"iii  its  pla.'e  .if  pni.Iu.'ti..n  (..r  al.siup 


I  l<lr    V»  III    AM>  TIM     ri  IIIM     lUdHIS 


fiJ'J 


tiling  to  iIm'  l<iilin'>H.  Tlii»''  iiroliliiiiH  linvi'  i-ii'i'iitlv  liccii  very  cnii,;.!!'!'- 
nlily  siMi|ilili<'i|  \i\  \hv  i '.  iMir.ilinii  "f  tin  iiriMiniti-  iiiftliiMl  I'or  tlic  oliiiia- 
tioii  III"  //'<    nrli-iii  (/  iiiiih  III  III   liliiiiil. 

Ily  iilix  I  \  iiiir  cliiiiiL'i's  in  till'  iiiiiHiiiit  iif  mil'  iiciil  in  tin'  MimmI  nitlicr 
tliiiii  in  llif  iiriiH'.  the  i'x<Tc1(iry  I'm-tur  is  piiillv  iMintnillnl,  iiml  it  I'iin 
Jic  ciiiiiiijcti'ly  Ml  if  iii'iiK'  ;iiii|  liliind  .-in'  ImiIIi  in\  f -liu'iitnl.  TIimiiUh  t" 
till'  wink  111'  I'l.lin.  it   is  iiiiw    |ins>ililc  tu  ili'lfi'iiiiiit'  witli  nil  "\1lciili'  ili' 

(.'I'l'i'  III'  jii'i'iinii'y  till'  mil-  iii'iil  in  tis  litllr  ;is  In  i- v.  >>(  hi I      Tin'  iiii|iiii'- 

tiilii'i'  'il'  lliin  Jirliii'\  I'linnt  will  lir  Miiprcfiiilril  \\  Inii  wi'  >tiiti'  tliiit  inior 
til  Fiilin's  Win  I;  nn  irnliKMl  r\isli'i|  liy  wliii'li  iiiii'  iiriil  cuiilil  Im'  .ippnix- 
iinati'ly  iiiriiMiicil  i'M'Ii    h  Inn   linui-  i|n;inlitii'     nt'   liluml   were  aviiiliililt'. 

Miirii  III"  tl.i'  work  tluit   lias  litiui  iliinn  hy  tin'  iisr  III"  lliis  inw  iiii'tlmil 

lias  sii   far  ii|i|>liri|  tu  tlir  ainniint   nt"  iiiic  m'll   in   ll"'  l>l 1  "t"  man   in 

\aiiiiiis  (lisrasi's.  \Vi'  sliall  ri'lVr  to  tin'si'  results  iiiiini'iiiati'ly,  liiit 
iMi'aiiw  liili-    it    is    iiiiiiintaiil    to   i-all    altriitimi    to    sunn'    wvy    simuji'stivi' 

ipliscrvatinlls    (•iilli-iTnillL'     ll'i     niiiililinil     nf     lirir    mill     in     lln      hlninl.        For 

iiiaii.x'  yi'iiis  tlnii'  liaM-  lirrii  in\  rsi  iu'atni  s  \Oni  liaxi'  tliiiii'_'lit  tliat  niii' 
ai-iil  I'Mii  not  III'  siiiiply  dissoKril  iu  lln'  IpIihmI  plasma,  liko  siii/ar  m'  sniiii' 
iiior^ranic  s:ilt.  It  is  liclii'Vi'il  liy  many  that  at  h-ast  a  pnrtiini  of  tin"  iirif 
ai'iil  riii'iilati's  ill  nMiihliiMliiiii  willi  niii'Icic  itliyinii-i  ai-iil  i  si'c  p.n.'i'  (i:!T  . 
wliirli  woiilij  ai'i'iiiuit  I'lir  tin'  I'ai't  lliat  siniii'  pniiiics  arc  calaholi/.i'il  in 
till'  hii(l\-  wlii'ii  tlii'N  arc  <_'i\i'ii  in  a  romhiin'il  state,  as  tliyniii'  ai'iil.  hut 
arc  cxcrcti'il  iiiicliaiiircil  when  int'cstcil  in  a  I'rcc  state.  When  jriven  freely, 
••erlaiii  pnrincs  aih  nil!' .  I'm'  cN.-miple  ma\  iiiurcnx  er  cause  inf!;:mma 
tiiiii  ami  caleiilns  riiiniatlin,  in  the  Ui.ii,i\s  of  ilnirs.  a  result  nm  nhtainci! 
w  lien  thymic  aciil  is  feil. 

Other  iihsciM'is  ha\e  <  nnelinleil  th.it  uric  aeiil  exists  .is  two  isnmeric 
varieties,  l;ic1aiii  ami  hictim.  the  mnmisiMlinm  sjilts  ni"  which  are  of  Uii- 
cipial    stahility.      The    less   stahle   u-snlt    is   much    more   soluhle    in    Mond 

scrum  than  the  stahle  /isalt.     ll   is  the  a-salt   that  In iiies  increased  in 

the  lilood  in  yolit,  the  dcpiisitiim  of  urates  in  the  tissues,  which  is  the 
most  characteristic  symptipiii  of  this  disease,  hein'_'  caused  Ii\'  convoi'sioii 
of  the  as.-ilt:;  into  /isalts.  The  structural  foiiiiiilas  of  the  two  isiunors 
are  as  follow  s: 


It 


UN  r  :  ()  N  rmi 

I 

O  :  ('  (•     Ml  lili.c  <■     MI 

I  ^             \  I  I             \ 

I     I            CO  ]  I            CO': 

I  -i         /  :  !         ^ 

II. \  (•     MI  N  '•     N 

I  l;iil;nii  ni".lnic;iti"ii  li.n.iiiiu  i  hiclini  iiin.jiiMnl  i  i|i  l'":iiiii., 


U 


ll>t:iiilt'   a  lli:itr--| 
■lalhi'lv  w'liil'lr  I 


still. ll'  ,-i  'ii:iti'^l 

( i'rljtti\i-ly    in^'tlulilt') 


'Pn 

n 

mw 

"  :      1 

1         ■ 
'l  ■ 

•^ 

1  •■ ' 

i'li 


i 

■  i 

;  i'i 

'  ■( 

y\\ 

iW' 


i       1 


M  I 


!      '! 


i        ¥ 


ii.-l' 


!^  r 


:i  . ' 


111 


I     1 


li 
11 


G50 


mi:t.\ii(ii.ism 


The  iiic.st  ici'.'iit  work  of  S.  I{.  Mincilict   Ikis  sIkavii  tliat  uric  jK-id  ex 
ists,   cliiclly    in   .•omliiiiiitinii    in    tin-   lilood    of   most    iii:miiii;ils   hut    not    in 
tli;it   of  th,.  Iiinl.     It   u;is  fouii.l,  for  .•x;ini|>lc\  tli.it    frcsli  ox-Mo«.l  ,'\,iiii 
iiic.l  liy  tlif  I'oliii  iiii'tli<»l  coiitiiiiis  only  i)M(»\'t  om.  free  uric  ,ici<l  per  Inc 

-'"■    "''  I'' 1:   :il''<'''   l>oiliM«r  till-    urolcjn-frcc    Mood    (iltrntc    witli    livdi-.. 

cliloiic  ;i.-i,|,  li,,NM.\,.r.  til.'  uii.'  jK-i.l  incicMscd  by  ;d)()Ut  ten  times.  Tlii. 
liiru'cr  ;iiiioiint  \mis  mIso  f.nmd  |. resent  in  whole  hh.od  tli;if  h;id  herh 
jillowe.l  to  staii.l  for  siniie  time.  in.lie;itini;  tliiit  the  urie-iicid  comi|>oiiihI 
(Mil    he   split    hy    iile.ins   of  ;in    en/.ynie.      The   eompouild    exists    in    the   eoi 

piisri,..  i,nd  not   in  the  phisnm.     It   is  of  s. si^rnifieiinec  tluit  nffer  tlm- 

settinir   five   )h,.   uiie   :i,-id.   there   should   he   .'ihout    oO  per  cent   more   of  il 
present  in  th.'  IiLmmI  .if  th.'  ..\  th;in  in  tluit  .d"  th.'  hir.I.  wliere  most  exi>t. 
HI  a   fr.'e  slut.'  in  th.'  serum.  iilthou-_di  th.'  iiruie  .d'  tlii'  ox  c..iitiiiiis  oiih 
the  sniall.'st    tra.-e  ..f  uric  a.'i.l,  an.l   that    of  th,'   hir.i   is  load.'d   with    it 
Iii\i'sti._Mli,,n  ..f  th.'  .• liti.Mi  .d"  iiri.'  a.-i.l  in  human  hlo.i.l  is  at  in-.'s.'iit 


III  lu'.iiri'css. 


Uricemia  in  Gout  and  Nephritis 

Th.'  prai'tical  appli.-ation  of  llies,'  ohsi'rvati.iiis  is  iiarticularly  iiiip..i 
laiit  ill  .•..mi.'cti.ni  with  //-,  .llnl,,,/,/  „(  ,,„i,l.     In  typi.-al  cas.'s  of  tliis  ,Iiv 

.■as.',  th.'  mi.-  aei.l  .d'  th.'  hi.....]   in.-r.'ases  fmm   its  iDrmal   vali f  I   t.. 

.'!  nur.  ii.'i'  ('.'lit  t.i  nearly  10  mi.'.,  in.li.'atiiu:  a  c.insi.lerahle  dcfrree  ..I 
r.'iial  iiisul'fi.'ii'ii.-v.     This  uri.-.'iiiia  can  not  in  its.'lf,  h.iw.'ver.  he  the  can-.' 

id"  111.'  .lepnsiti..ii  ..f  nrat.'s  in  th.'  .ioiiits,  h.'.-aiise  it  ah, curs  in  ..tin  r 

.Iisea>.'s  with  r.'iial  reti'iiti.in.  sii.di  as  n.'phritis.  M.n.'.iv.'i',  the  hl.i.id 
s.riini  IS  .-apahl.'  .if  -lissoK  iiej-  niiK-li  larjr.'r  ipiautities  .if  uric  aei.l  than 
ai-e  ever  f.iiin.l  pr.s.'nt  in  it  in  „.Mit.     The  real  i-ause  f.U'  the  L'.-Uty  d.'p.isit- 

must  depi'li.l  .III   s.ii hail.;.'  alT.'.-tiny:  til.'  hl.i.i.l  s.i  as  to  all.'r  th.'  forin 

III  wlii.'h  uii.'  a.'id  .'xists  th.'r.iii,  with  the  r.'siilt  that  it  is  excreted  int.. 
the  Joints  an.l  deposit.'.!   there, 

Othi'i'  dis.'as.'s  slniwiiiL'  uri.-.'iiiia  ar.'  Iimf  innxoiihuj  ami  tu  plirilia.  In 
the  latt.'r  disease  the  dama'_'ed  cx.'r.'t.uy  fiin.-ti.iu  of  the  ki.lney  i~ 
iiianil'.'st.'.l    first    .if  all    hy    an    in. Teas.'    in    the    iirii' a.-i.l    c. int. 'lit    .d'  tli. 

'''"'"'•   ■■ 'iii|.ani.'.l   later  hy  a    r.'tenti.in  of  ur.'a   an.l   later  still   hy   .hm 

of  .'r.atinin.'.  Th.'  s.'v.rit>  ,.f  the  renal  iiiv.dv  I'lri.'iit  may  therefore  h.' 
■.'aiii!-.'.!    li\    il.t.'iiiiiniri.j:   th.'   p.'reeiita..'c  of  these   three  in.'talxdites.      On 

•■"''■"""'   "''  'II''  iin[i.'rlai .if  these   facts   fr.uii  a   I'liiii.  al  st.au.lp.iiul,  \\.' 

app.iid  a  tahh'  .■.intainiii!,'-  results  s.'.-iirc.l  hy  Myers  an.!  l-'ine,  in  wlii.-f 
th."  h.'havi.ir  of  th.'  m.'taholiles  in  t!ie  hhi.i.l  Is  slio-.  in  rehdiohsliii- 
1.1  the  severity  of  th.'  case  as  irail^'i'd  hy  the  hlood  pre     are. 


ruic  Acih  AMI  Till;  ituini;  I'.oniKS 


651 


I'r:rc  Ariii.  l'i:iA  N  ami  ('i:i  aiimm.  m   Ulcdh  in  (lui  r  ami  I:ai;i,y  axi>  Latk  Xki'IIRITIS 


I  i;ii' 
Aim 


1  i;i;a  n        i  kkatimnk      systolic 

III.OOI) 
I'KKSSrilK 


Mli.    lo    lIKI   I.e. 
Ml,i«i|> 


T\|iir:il    Cli-is    111'    I 


Tv|iir;il  li;iily  Iiilrr>  ■   'ii:  Vrilu'lis 


Clinniii'     DilTii^..'    iin.l    Chnniic-     liiln 
stitiiil    Xi'j']iiitis 


N.I 


.1.0 
7.1 


t»..'l 
11,.'! 


I.'.t 


1 


17 
1  I 


li'. 
ix 


•2\ 
i  I 
lit 
f'i7 
17 


*'..) 

•J  J 

l.-,.ll 

-1(1 

1 1.:; 

mi.". 

i:;.ii 

■Ill 

S.7 

111 

1.1 

■SMI 

L*.- 

nil 

L'.t 
1   7 

i;oi) 

1 .  1 

1  v-> 

•'  * 

i.'-iii 

::.!» 

i:;u 

iV't 

■>  1 

1S5 

100 

>>  1 

U7 

l.-)0 

l.s 

L'HI 

l^.'.i 

170 

I.'.i 

1 1.-) 

I.'.I 

L'lO 

l:;o 

'■'•A 
1.(1 

lil.-j 

.l.O 

L'on 

111. 7 

LMO 

'           !.''.■• 

--■' 

111*.- 

li-O 

11.1 

ijiiri 

11.0 

■fvpi.Ml    l-'riliil   Cliiiiiii..    Iiili'islitinl 
Ni'i'hi  ili.H 


(Mveis   .'III. I     Fiiir:       Auli.    Int.    Mi'il.,    lUl".) 

Liistly,  i-cuMnliiiu:  tlic  iiilliicMiT  nl'  ilnii;s  mi  tlir  lilm"!  iirir  iifid  in  ilis- 
i';\sc,  it  liiis  li(>(Mi  I'omiil  l>v  I'iiic  tliat  linlli  ;itii|>liiiii  Mini  siilii-v  Intos  caii.'ic 
:i  iiriiiiniincoi]  (Ici-rcjisc  in  tiic  ariiuiiiit.  loit  tliat  it  i;i'!i(liially  rises  to  llic 
iiiil  li'vcl  oM'ii  uliili'  ailiiiiiiistratiim  nf  the  iIiul's  is  lioiiiL'  cnutiiiuiMl. 

liiipiiiMaiit    <-iiiiliiliiiliiiiis   to    tlic    liclia\  ior   (if   I'i'ic    acid    in    !•! 1   iivc 

cniistaiitlx'  appcariiiu'  at  |iicsoiit.  inaiiiiv'  fniiii  tlic  lalioratdrics  (it'  Foliii 
in  l?(isliin.  Ill'  S.  !{.   Mciicilict.  and  of  .Mvcis  and   Fine  in   New   Vork. 


'= 


jib 


I  s 


ippl 


(I  : ;( 


ilii 


[f''^ 


i        1: 


'  1 

'■      i 

^ 

1 

p 

I 

lltliS, 

1 

(•iiAi'Ti;i;  i.xxiv 

TIII-:   MIlTAI'.iil.isM   or  TiiK  t  AI.M'.dll VDK'ATKS 

Tlic    Iirallliy    ;iiiiMi;il    ,ir-,-iniMii    is    .•;ip;iM,.    of    r;il>i(lly    oxidi/in-    liir.j. 
-lUiHltilirs    or   .•;,,I„,l,v,ii;,|,.,    ,:s    is    ,.\i,l,.„l     fV.M.l    tlu'    i'.iliowilljr    f.-iHs:     "|  I 

.■.•irliMliy,!!;,),.  i>  Mivrii  1,,  ,,  st;n-viii-  .-iiiiMi;!  I,  (  li   (lie  nuTirv  .mtimt   v.t\ 

sliurtiv  ;,rtr,,,;,r,I  in,.,v,-,srs:  •_>  .  tj,,.  ...spinitnry  <|n,,1ir.||  ills.,  in(T..,Ms.s 
in-Il.Mllllii   ll,;it.   .vlMtiN,.ly   t.,  MSy,.,,,    inlllk...   „„„•,.  .-iirlM,,!   .lioxi.l,.   is  Imm,,, 

<'x.-iv-,.,|    IS,.,.  ,,|,.r,,  ,;J7,.   ,,,„|     ,;; ,.    ,1^^,    j,,^,,.^,,,,,    ,..,,i„,|,v,|r,K,. 

'"■■''''■•^   ''-^  .•i|.pc.-ir,-M in   tl„.   ,.x,-n.t,'i.      In.lr,.,].   „f  ti,,.   tluvo   ].r..xiin,-it, 

IMinripirs  nf  Wuu],  ,■.•,  ,l„,l,y,Ir,-|t,"  is  tli,.  „„,st  avjiilahlo  for  (M.nilMlsti,,!, 
'"  ""■  ""'"'■''  '""1.^-  "  "uy  tl„.n.f„n.  I„.  ,-,,nsi,I,.|v,l  :is  tlio  ,|ui,.Ki; 
ii\;iil:ililc   I'll,'!    |',,r  ili,.  l„„]y    rin-iiiiccs. 

CAPACITY   OF   THE  BODY   TO   ASSIMILATE   CARBOHYDRATES 

Assimilation  Limits.     \Vli,.„  il„.  liniit  t,,  t!„.  .-nnnunt  of  .•.•irlmliv,],:.!,- 

"''■"    " '■-•■iiiiMi.  .-.■in   inrl.ii  ,,|i„.  is  ov,.rsf..i,!.,.,l.  soni.-  of  it   iipptMrs  ii 

t\u'  uriiMv     Til,.  .-inhMu,!  il,,,,  ..,•,„  1,,.  1ol,.rat,..l  «itlio„t  .•.•uisiii.,'  .Jv iii., 

IN  coiriinoniy  rail,.,!  tl„.  „.„;„,,/„/;„„  ,„.  ...„/,„■,,//„„  i;,„;t.  Tlu-  usi-  .,f  ti„. 
fcnn  '•linnf   is.   !„,„,. v.., ■.   v.ry   unforl  iiiiat.'.    ;•,„•  it    i,„pli,.s  tlwit   ]H-y.,u<\ 

""■^   ' '    ""■   "'•■■^■"M-iM    i-   .-.ip.-iMc   of  .i,.,-ili,;.^'   „it|,    no   „„,r,.   ,..-n-liol,v 

'''■'""•  "'"'•'•  '^  '■•■"■  '■'■ '"i"-  11"'  '-.I-'.  I'or  if  M  j.-n-,.,-  aiMount   is  fak,.'i, 

"'"'>  ■•'  ■^'"•■''1  "■^"•''  '"■  '!"■  -N--  "ill  .■,pp,.,n-  in  tl,,.  in-in,..  Wli.  n  1l,. 
nnn-    ,s   .•,ll„„,.,l    to   ..„!!, .,.|    f„,    i  „  ,.ntv    f,,,,,-   Innirs.    tl„.    mixr,]    spn-i„„.;i 

^'"'"~-  ""  "■■ "'■  ^'"'-"N''   in    111.'   n,n.i,,rity   ,,f  liraltliv    ii.,livi,lii,-ils  aflrr 

til.'  ii,....sl,„n  of  L'liu  ,„„.:  ,n,.,.  .-KM)  .,„,,  ,.,  son„.„li,,t  liiyluT  ix.m.nfaL- 
"I  ■•••i-.'-^  .I.n,.|,,p  a  mil,!  .Jvosiiiia.  ImiI    fi-.,pi,.nlly  i„,iu>  is  ovi.l..nt  .'v,.!, 

:'"'■'■  •""'  -'"■     '■"■■^ I  'I"'  I'-'^l   iM.'iiti.ih..,]  ani..iiiits  111,,  limit  of  ii„_.,.sti,,ii 

'■"   '•'■■•"•'"■'I.  ""    ■' lit    of  naiis,.a.  ,.t,...  ami    it    is   iinpn.l.ahl,.   tliat     .■v,- 

"    '"'■-'■'■  ■■ '""-^  '-""I'l   I-   to|,.rat..,l.  anv    .„.,.•.'  of  fl,,.  .l,.xtros,.  uoiiM 

!..■  ai.sorh.,1  than  uitli  ;l(l()  ,„■  4(10  .„,,  Tl...  t.^slinn.  of  tl„.  so.,-all,,i 
assiMiilatioi,  li,„„  l,,s  |„.,„  ,.„„si,|..,v,l  ,,i  important  ni.l  in  llu  ,Hw,n„<,. 
of  cnrl,,  r„s,.  ,„  ,h„h,l,,,  ,1„.  ,.|,ara,.t,.,ist i.-  f..Mt,nv  of  s,,,-!,  .-as-^s  lioii,.. 
""'  I'l.alnhty  ,,f  tlir  ,,i-.janiMii  t,,  assi,„ih,|,.  p,.„p,.,iy  tin-  usual  ,|uantitv 
of  farlioliy,!!-,!!,.   i-,,nt,iiii,.,i    in    tin'  .lii'l. 

!!        lias       OCI-O         fi;!::;,l        ll.-.j         I,,        |.    .       il  ,  1,  ,.  ^ 

'  '  '    ■'■■"    ' '"     'ii<-    Hsnils    oi    anv    value.    ccMtai!! 

'■'""'" "  "'"^'   '"■  '"Hill''!  in  applying  ||,..  assiniilati.ui  t.-sl.     Tli,>  nmo 


Tin:  Mi'iAnni.isM  di-   riii.  «  aiji'.hiiyiiuatis 


(;.-:'. 


im|)(irl.    it   of  tlifsc  coiici'i-iis  llic  ;irti\  itics  iif  tin'   '_'.i>t  iniiitcstiii.il   nppn- 

I'iitiis  ;it   tlic  tiiMi'  till"  siijiiir  is  uiv.'ii,  for  it   li;is  I ii   \'<>\\\[<[  that   it'  (itlii  r 

fiMKlstulTs  jirc  Ipciiii;  alis(pilii'(l  at  tin'  saiin'  time  as  tlic  suirar,  nuifi'  ui' 
tin*  latliT  call  he  tulcratcil  tliiiii  wln'ii  llic  siit;ar  aloin'  is  lu'iiiir  alisdi-iit'il. 
It  lias  tliiTcfori'  lii'iii  iMisti  iiiar\  tn  'jixc  tlic  siiuar  ili--,,lvcii  in  water. 
nr  in  weak  cnlTcc.  llic  tiist  tliiipj  in  the  iinn-ninii' after  the  |iaticnt  awakes: 
i.e..  at  Iciisf  twelve  to  sixteen  lioiiis  after  llie  lasl  nnal  was  taki'ii.  In 
iiiaUiiiy:  tlicsc  tests  tlic  mine  Ndiilc.l  liefcnc  the  siii^a''  is  I'stiinated  slioiiM 
of  course   itself   lie   tlioroiielil v    c\aiiiiiieil    for    lediieine'   siilislaiices,    ami 

tlic  urine  slioiild  lie  (■(illeetecl   e\  er.\    ninety   iniiiiites  and  cxatiii 1   \'y  ii 

relialilc  test    (  licneilic-t  "s  or  N\  lander's  >  .* 

Alllioiiu'li  a  limit  is  set  to  llie  ahility  of  llie  oi'vanism  \\<r  retainiii!.' 
sugar  iiiioiio.  CM'  di-saceliarides  I ,  this  does  not  seem  to  apply,  in  liealtliy 
indivi<luals  at  least.  mIicii  starches  '  polysacid'iarides  -  are  inuested.  Tliils. 
it  is  a  welhknown  fact  tlia;  jieopli'  can  eat  eii<n'iiioiis  (|nantilies  cd'  pota- 
toes or  of  lii'cad  without   the  appearance  id'  aii\    tra f  redncinir  siih- 

staiici's  ill  the  twenty-four  hour  urine,    dii  the  (,ther  hand,  uii ilh'clcd 

and   csainiiicd    at    short    iiitcr\als    'cxeiy   half   hoiiri    after   takinu'   lari.'e 

(plantitics   of    polysaeellaride-rieh     I'nnil    will     I'lviplelltl  .     he    fnlllld    to    eolltaill 

traces  of  rcdueiiii.'  sulistaiH'cs. 

I-'or  i>ractical  inirposes  it  has  hceii  cuiisidcred  that  an  ind  ln;d  who 
develops  ulyeosuria  after  takiii-  Ino  em.  of  ejiieose  must  he  ;isidcrc(l 
as  at  least  a  potential  <lialietic.  In  the  lieht  of  the  almve  results  and 
foi-  many  other  ri'asoiis.  there  is,  howcM-r.  coiisiderahle  doiilit  as  to  the 
value  of  tlii>  assiiiiilalinn  test.  Thus,  when  a  sulutioii  ■  '  ehi,.,,se  is 
Ldvcii  orally,  its  rate  of  alisoiplioii  will  depend  \ciy  'aruely  on  the 
iiiolility  of  the  stomach,  if  this  is  i;oiiiial.  the  solution  will  very  quickly 
find  its  way  l>ast  (lie  p,\  lorie  sphincter  into  the  intestine,  where  it  will 
ho  rapidly  ahsorhed.  If.  <iii  the  other  hand,  the  pyhu'ic  spliinctcr  docs 
not  open  freely,  the  i>ass;ii;e  of  the  ehn'ose  into  the  intestine  may  lie 
so  delayed  that  no  more  is  inesenl  in  this  i)liiee  at  one  t'liii'  than  would 
ho  the  case  after  an  orilinary  diet  of  polysaccharide.  .Vml  even  after 
the  siiirar  solution  enters  the  small    iniestine.  dil'I'erciiccs  in   the  amount 

of  the  intestinal  contents  with  which  it   hi mes  mixed,  in  the  extent  of 

hactcrial  irrowth,  and  in  the  ahsmption  pro' ess.  may  very  materially 
affect  the  rate  at  which  the  uhii-osc  e;iins  mitry  to  the  hlood. 

AltlioUL'li  often  of  douhtfiil  diaunostic  \ahie.  dcteriiMiiat ioii  of  the 
a.ssiiuilatiou   limit    is  of  coiisiderahle   aid    ///   i-diilntlliiiii   tin    tniilniiiil   uf 

•Examin.ilion  of  iuirni.il  ituiivijiials  lias  .-Invvn  tli.it  tin-  i~~innl.Tti'"ii  In;. it  i't  .lilTiniil  Mn;:ir^ 
v.iri.-  -mir  rtlat  ;  (or  bIiu-..-.'  il  :iM-.ii-  1..  1"  (i..!ii  ;ilM.m  I  I  1..  J-ii  u'li  ;  |..i  l.iiil..-,-.  «lii.ti.  il 
tt-:!l    he    rrTTirtnl-.rrnl.    U  thr  r:i-.nn=-:ir.-!!.ir!:l»*  .is-^rri.l!e(!    wi'll     'liv.-*--'    !•'.   tile   f nn«trit.-tt..n    nf   fhc    (■.it". 

I  iLir-'^e    it<o! 

h.ttl.T 


siikMr  niolcciilc.   the   .issii'iihiti'iTi 


till-  Ii. 


limit   il  fri 


1111!     In     1=11     „,■•    ;     f, 


I-    i.iiir-    •.itu.ir   i.r   s.ti 


i.l.J.il.lv.    l.'it 


.;i\  til    .-I-    1.  I\' 


1,1      ■'.,! 


f.ir   l.i>l" 


(lis.icili.irii!f.  .liiil  llu-  vuLMT  i<r.  sent  in  liulk.  tin-  a--lMii 


111.  11  Iniiit  i.  .li-tiiully  l.'Wir 


!  m 


jr       » 


!il 


m     i 


If 


i ". 


s; 

i  ■ 

i 

.1  j 

i-; 

'i  ■■ 

i'  11,  : 


U 


r.r.4 


Mi;T.ii!()i,is>t 


,ll,ih,l,.:      V„r  this  pnri.oM.  tli.'   palirnt  slir.iil.l   first   «{  u,-   instni,;,  i 

l(.  fi<]\,i\v  ills  iisii.il  .lift.  s..  Ili.it.  I.y  .■xjiiriiiiatii.ii  of  the  iiinoiiiil   of  si|.j 
-■MT.'t.'.l  ill  tlic  miih'.  ,-in  (>|>iiii..n  may  Im'  l'<iriii.'c|  of  tlic  s.^.Mity  oi'  i! 
i-;isc.     Till"  (lii't  slioiiM  til. '11  li.'  :'li,iiin,.,l  s„  as  t.i  consist  (,f  a   i»;nt   ll:, 
•  •i.iitaiiis    no    (Mil.oliy,|iat<'s   ami    aiiolli,.r    ..ompos..,!    .•ntiiviy    of   star. I 
l'"'..l.     Tlif   foniicr   is  ma-lc   up  of  .-vus,   lisli.   u-ivfii   v.-ctalilcs,   fat.  ..(.■ 
Mii.I  tlif  latter,  to  start  \\itli.  s|i,„il.|  consist  of  10(1  iiraiiis  of  l.rcad.  ,11, 
tnlnit.'cl  lictuccn  tlic  two  main  meals  (,f  tlic  .lay.  ..nc  „(  wliicli  is  lir.aK 
fast.     This  ,|ict  slioiil.l  l,c   ...iitinn.vl  until  tli..  -ly.-osuria  .'itlicr  ilisappe;. 

'"■  ••'"••li"'^  •• nstant   l..\,.|.      If  it   .lisapp.'ars,   tli,-  .-asc   is  .-lassiti..,!  ,k  ,, 

nul.l  ,„u-  ..f  .liali.'t.'s.  an. I  tli.- .laily  .■ill,.u  aii.'.^  of  l.rca.l  may  1...  iii.-ivav,  ,1 
l>y  r.d  uranis  a  .lay.  until  tlic  suirar  auaiii  mak.-s  i's  apii.'araii.'c  in  ll,. 
"I'iii.".  in.Iicatin^r  that  Ih,.  assimilalh.M  limit  has  l.ccii  icachc.l.  I',  - 
tlii'rai>eiiii,-  ,,iirpos..s,  tli.'  palii'iii  sli..iil,l  now  l.c  instnn-t.'.l  t,.  take  ali..ii' 
thri.e  t'..iirtlis  of  this  aiii,.iiiit  <.f  .•arh.iliy.lral.-  in  liis  .laily  rations,  am! 
he  sli,.ul,|  ]„■  siippli,..!  with  .'xprK-it  iiistrii<-ti..iis  in  Ih.'  sliai>o  of  .11.' 
tahl.'s  as  to  what  v.-iri.'ly  an.l  .piaiititi.'s  of  the  various  .-arlMiln  .li  at. 
mat.  rials  his  to.,,!  may  .•..ntaiii.  His  urine  sh.nil.I  he  exaiiiin,.,!  at  IV. 
<|ii.^nt    int.'rvals     ,,iic,.   a    u..,.k     an. I    he   sh.nil.l    l„.    iiist  ru.-t.'.l   as   to    li. 

"='""' '  '''-  <y>^<--^-^<'  Mil. I   the   importan,.,.  .,f  his   re„iainiiitr  au'ly.-osiiri.- 

I'.y    further   tivatnu'iit    such    so  ..alle.!    Ial..|il    easis    .,f   .liah.'tcs    iiiav    f. 
k('i)t   in    perf.'.'t   h.'alth    tor  many   ye;irs. 

WIicii,   oil    th.'   ..ther   haii.l.   th,'    .^ly.'.suria    .^xlsts    with    1(1(1   •■r.iiiis    .  ■ 
lir.'a.l  ill  th.>  , laily  ralhai,  this  must  h,.  r...|ii,.,',|  t.,  :,{)  u-rams,  an.l  if  aft. 
s.mie  .la\s  tli.'   tirst    re.lucti.ni   does  not   sufli.-i-   to   r.'ii.l.^r  the  iiriii.'    fr. - 
from    su-ar.   carl,..liy.Irates   must    he    «itlih..|.(    .■ii(ir..|y    fr.mi    tli.'    .li,  • 
ll    the   e|\<-.,suria   .lo.'s  n.it   n.iw  .lisappi'ar.   tin-  .-as.'   is  t..  he  .■onsi.l.M  ,■  ! 

■""■"■'■  !ii"l   i'   "lay   ]h-  -cssary   t.,  un.lertake   tli..  starvati.iu   treatiii..!  • 

\\lii.'h  has  re.-..iitly  l.<'..ii  .lev."l.i|....|  in  this  c.,uiitry  hy  .Mlcii'^  an.l  .loslii, 
with    api>aiviit    sn.'....vs,      |!y    th.'    re.liicti.m    .,f   cai  !...hy.lrat.-.    .,r   I.y    tl,. 
sfarvati.Mi    tre;itm..iit.    it    is   usually    p.,ssil.lo   t..    make   even    the   s.^u'r, -• 
.-Ms.^s   of   .|i;il,.-tes   airl \  .'..siii  i.',    an.l    when    this    lias   Iie.'n    attain. ■.!.    th. 
trra.liially  1..  iiu'i-eas..  th.-  aiii.,ui,t   of  pr.,t..iii   .,r  .'arlioliv  .Irate   f.io.l   iii.i 
the  assimilati.i.i  limit  iijis  ln'cn  rca.-li.'.l. 

Saturation  Limits.     T,.  av,,i.l  .rmr  caiis,.,|  |,y  irr.'-.ilar  ahsor|.li.,ii  fr... 

the  intestines,  s.,m.>   iiiv.'st  ii;-at..rs  liav.>   r .iiimeii.l..,!   the  .h'terminati. 

of  the  assimilati.m  limit  aft.'r  intra v.-ii.. us  ..r  sulH.|itaii.-oiis  iii.i....ti..' 
of  suu'ar.  Mut  ev..ii  this  rcfiii.'ment  in  t...-liiii.-  has  ii..t.  as  a  rule,  lia.l  tl 
•■ff.'ct  of  ren.h.riiiL'  the  irsults  „(  any  v.'i-y  ..vi.l..iit  valii..  as  a  "erit..ri.. 
of  the  utilization  .,f  ylucose  in  the  animal  h.,.ly.  The  roas..n  \\,i-  tl, 
iitir.lial.ilil>  ..r  Ih.-  iiietho.l  is  mainly  that  the  p.-ri...!  of  iu.jeclioii  of  th 
frliicos..  s..luti.,n  usually  ...•eiipies  only  a    few   minutes,  so   that    it   can-- 


THK    MI;TAU()I,IS.M    of   Till:    CAUlidllVliKATliS 


655 


a  snddoTi  instead  of  a  vciy  (;'""liial  iiiiMf.isc  in  tlic  sujrar  ciinccntrafion 
(if  till'  liliiiid,  till'  i'i)ii(litii)ns  lii'iiiu'  quill'  iiiiiilu'  tliosc  wliicli  i-xist  iluiiiit; 
till'  iimiiial  alisorptiiin  of  i^liii'osi'  fioni  tlii"  intrstini',  Tlu'  iiiccliaiiisin 
liy  whii'li  the  Imdy  ordinarily  disposis  of  excessive  a iints  of  glucose 


alisorlied  into  the  portal  Mood, 


)t  ad 


justed  to  operate  when  the  sys- 


teniii" 


hlood   is  siiddeiih-   ovei  eliarj.'ed   \\ith    th 


ihsti 


In  the  one 


is  a    foodstillT;    in    the   other,   lieeaiise   of   its   exeessivo 


ease   the   U'llleose 

ponei'iitration  in  the  Mood,  it  is  more  or  less  of  a  pois  iii.  Such  results,  in 
otlier  words,  meri'ly  show  lis  how  iiineh  udiiense  can  he  acMed  at  one 
time  to  the  oifiaiiism  without  an.\  oveillow  into  the  urine,  hiit  they 
furnish  us  with  no  inforniation  reirardini,'  the  |)ov\er  of  the  or(;anisin  to 
•itili/e  a  eoiistant  thoiiirh  moderate  exeess  of  this  siilistaiiee.  In  the  one 
ease  it  is  the  "saturntion  limit,"  in  the  other  the  •"  nlili/.ation  limit"  of 
the  ortianism   for  irlueose.   that    we  are   really  eonsidei  iiij;. 

Consideration  of  these  ])iineiples  has  led  Woodyatt.  Sansimi  and  Wil- 
der-" to  uiidertaUe  a  thoroiij^h  reinvest ijrat ion  of  the  whole  i>roMeiii  of 
the  utilization  or,  as  they  prefer  to  eall  it,  tin  fiili  niiiii  (if  tin  liixlii  jiir 
i/lurose.  Tliey  empliasi/.e  the  ohvioiis  fact  that  the  aliility  of  the  ortranisin 
to  utilize  tilueose  "iiiust  depend  on  the  rate  at  which  the  tissues  are 
ahle  to  alistraet  it  from  the  Mood  hy  their  eomliined  powers,  to  luirii  it, 
to  reduce  it  into  fat  or  to  polymerize  it  into  ud,\co!,'en  "  To  form  any 
estimate  of  the  eonihiiied  effect  of  these  pro<'csses.  wc  must  take  into 
account  not  only  the  amount  of  jilucose  jier  unit  of  hody  wei<,'ht  irrams 
(ter  kilotfram'),  liuf  also  the  rate  of  injection,  for  "tolerance  niust  he 
reirarded  as  a  velocity,  not  as  a  weight." 

Hrielly  siimninrized,  the  conehisioiis  whii'li  \Vood\att,  etc..  ha\e  so  far 
drawn  from  their  invcstifratioiis  are  as  follows:  In  a  normal  rahhit.  dojr. 
or  man,  OS  ().!•  frm.  of  glucose  per  kilo^jram  lioil\  wi'iirht  and  i)cr  hour  can 
he  utilized  t>y  the  or^ranisin  for  an  indeliiiite  time  without  causint:  jily- 
cosuria.  AVhcn  lietween  0.8  and  2  trin  fre  in.jccted.  a  i>art  of  the  excess 
api)ears  in  the  urine,  steadily  inereasiiij;  until  .i  niaximum  is  reached, 
after  which  the  excreted  fraction  remains  constant  (;it  ahout  oiietcnthV 
If  more  than  about  2  urams  per  kilofjram  an  hour  are  iii.ji'cted,  "a  larire 
pereenta{»o  of  all  (jlueose  in  excess  of  the  '2  pm.  per  kiloi;r;im  an  hour 
appears  in  the  urine  ulipii  constant   conditions   are   once   estaldished. " 

The  fact  that  so  much  <;lucose  in.jccted  intravenously  can  he  used 
without  the  aiijicarance  of  any  of  it  in  the  urine,  indicates  a  method  hy 
which  foodstufTs  may  lie  sui>i»lied  to  the  tissues  in  cases  where,  on  account 
of  f»as1rointestiiial  disturliances,  it  is  im)iossihle  to  linve  food  ahsorhcil 
by  the  usun'  pathways.  The  possible  value  of  such  a  method  of  treat- 
ment in  cases  of  extreme  weakness  has  been  tested  on  laboratory  animals 
b.v  Allen,  who  states  that  such  iii.jcction  seems  to  have  a  valuable  mitri- 


i  i 


»!i 


il    H 


■  i 


M   I 


i 


k  i 


■h  » 


11 
■  1 1 


!■ 


I'Ml  'H 


:  si 

1 

t.  ■ 

\    ■ 
j    • 

1     ■  ^ 

? 
( 

« 

f 

■  l 

I 

t 

1    ; 

i 

!    I 

.  ! 

^^^^^^Uk  *. 

1 

I 

I'l 

U'  tt 

'  i 

'1 1! 


.  i'  <     ■"■ '         S     •  til 


:i ,  f 


i,N« 


(;:>(; 


.mi:t.\ii(ii.ism 


"•"'    ';';7   "•-'•;•   ''1 1    tnn,sf„sioM    is    in,pnH.,i,..I.lo.    .1, so    in 

Ms,„nssl,„„i,M,„  ,,,.„,,     ,,„,,, in,i,.,.,i„n  ,„  s„...n-.  ..1,1,..,.  f,„.  ,1„ 

"'""":"';'""■;■'" -  "'••  ■•'-i-'ii-'-n  n-Mi.  „..  ui.i,  ,i,.  „,„•„„,  „f  „,, 


s.m,..,i>M,.s  si,.„d,i„sr  iw.u.i,„.,..i  „,  ,1,..  ,,„i„t .,,.  i„j,.,:,i,.„ """  "'"' 

,,,)';;  J:;?,V'''^ '''•■'';;'';'' i'i"-''i'-f^ t,,.,,..,. r„.si,nii.. 

i"""   ;'"':"'"  '■ "•  ^■""•■^'  '^^■"  '-'>•  """■  ""•-,  .,,„. 

'■'■"'"  ''"'"'-   ""•"■  l'.-'-.-'^"'  'ImMml,  ,1,,  ,ni„„.,l   |„„U, 

DIGESTION  AND  ABSORPTION 

Dig^stion^     .\ll.li...s,„,,,.,„,,„,,,,.„..   „,.,,„,,   „,,, ,  ,^  

'/''":"•   '","'"'■''   "    -••'''-'■'"•'1    inf..  .!„.  1,1 1   „r,l,..   p„,,,.,l   svs„.„, 

'';'r  "'""'♦    "''^   '••-""■ i---''..V.l.....in.,.   ,„„n,.s.....l,;n.i: 

;;::';:;  ""■^'"■'■":";''"^'"'  •'■'"•  "-<"nws..  is  ,i,..  ;.„„;.  .,rs.iiv.: 

^t.'itr.   I.....   ii.,t   surnMin,],.,]   I,y   ;,   ,.,.ll„l„s,.   ,.nv,.l.,,„. 
i.-li.  P'.v,.|lm  .lisplays  liftl,.  of  its  .-...tivify  ,],iiii,.,r  (1,, 


"'•■i'    ■■II-'.    ill   a    IV,.,.  Stat,.,    i.  ,.., 
''lit    I'Vi.!,   III!    iV,.,.   stai-,- 


(    111,,    tl,..     !•  1     •        •         .1  ,  -    ..■    .  i\  II  ^      l|,iiril'_r     III'. 

"'";'.""'"'",""■• "'■     Al-.,.,.  ,!:..,• lissuall-,w,.,Ian,ll 

'"'.'"""';.';'"; '-'"■""-'■•— ii.  .1,,.,...  is  a,,  i,i,,.,.vai  ..r  ,i,„.. 

^-nn.  unt,ll.,,l,.,i..li,,,,i,.  ,,,,,,,,,,,,..,,,_,,,,,,,,,  ,_^^^^^^  ^^^^^  ^^^^^^'^J-^^ 

!:rr;; ;"•";'■--;'"--—•' ^'^  .inn,i..i,i,.i,n:.;t  , 

t     .,n.„.sta,-..l,,„  ,1.,.  ..,.,„„,,„,   , ,       „„,,„,  ,„is,i „„.,., i,i„ 

'"■'":^""r  '"••""">■  --"••' i.f  .1,.'   la,.Ml,a,    a   sli.!i:. 

""  '■■''^;i '"  ''.y''""""-' .'iitrati,,,,  „r  ,1,..  ,ii,„.„i,„  ,„;,„„,,.  _.„.;,, 

crad.s  til,.  ii,.ti,,ii  ,,f  ptyaliii. 

TiH.  ,,,.,.,,,„.,  „r,n„,n^,.,,i„„i^ „..,..,  ^,  ,,^ ,,^,,., 

o.,,W,..s  ,.,,,,.,,., .i.<-ii.i.,l,..i„t..s,in...„.....a,.l,.,,,v.i,.a.,.. 

''     *  "1  '•■"; '^^  "^  '"^" '  ^""•'•'--  l"-"'b   .''^  .'Inn.s,..  an.l  parth 

";  ""^;'      '"  ""■  l-"-''l-nvi,.'„„„..,i,  .,,• , ,,„„, „„  /„„„.: 

"  "";"';''  ''"V """  ""•'""-■    ''-'"•  •"••'1—  'i-iv...i  ri„,n  ,1, 

.nvl,  an.l     li..  ,ni,.l,an.,.,i  sn.ars.  s,,..],  as  .-an..  s„.ar.  nialt-.s,.  au.I  la- 
'""''   '•""•"■""•■•^"""    m   ,1,..   H.t..,,inal ,..n,s.   ar..   n',t    i„n iiat..lv   al. 


i-i .  !• 


!l  I 


Tin:   mi:t\iii>i.i>m   nc    iiii    i  akiiuiimii;  \ii;s 


li.u 


^iirlicd  info  tlir  lilninl,  lull  liiTiiliir  MiIp.JccI  In  llic  ilctiiili  nj'  nlliir  OMZyillcs 
.(iiilrilintol  li\  till"  iiitcsliM;il  jiiiiT  ii:iiiicl,\ ,  tin'  inn  rliii'i  iiiniiiiis,  inic 
of  wliicli  exists  I'm-  cai'Ii  ol'  llii'  <lis;ic-i'li;iii(|is.  I'>\-  tln'ir  :ic'tiiin  iinilt<)st' 
is  (•(Hivcrli'il  iiitii  twii  iiiiilfciilt's  lit'  "^hiccisc  li\  the  fu/v  iin'  iiialiiisc:  l;ic- 
tcisc.  into  ^'iil;ictiisc  Mini  ulllc'isi'  liy  liirliisi-.  iilhi  imiic  siij.Mr,  iiiln  j.^x  ii- 
liisc  iiinl  L'lni'iisc  |py  invcitiisc.     It   is  inti'i-csiin'.'  to  note  tlmi   in  .•luininls 

wlmsi-  t'i''"l  ildi's  i.iit  ciihliiin  oni'  o|'  tlins.'  dis; iniriili's.  llir  i-di'rrspunil- 

\\\\x  iii\  I'ltiii','  c'lizynic  is  iiliscnt  iVi'in  liic  inti'siiniil  .Itiicr,  Tiii'  lii'rliixn- 
iiius  ;ii:ini;ils,  I'm'  cxiiniiili'.  A"  imt  I;ik('  :iii\  lai'tnsc  In  tlu'ir  fiMnl.  and  tin' 
intestinal  juice  contains  iLciermr  no  lactase.  altlinni;li  it  is  jiicscnt  in 
tliat  id"  tlie  younu'  animals  wliile  still  siickliiiLr. 

A  certain  aiiinnnt  of  cailiolixdralc  liccnnn's  attad^ed  l'\-  tlw  mli  ■il mnl 
hdi-lii-iit.      Tliese   split    tlie    inniiosaci-liaridcs    into    Inwer    I'atty    acids    and 

'.•ases.  snidi  as  iih'IIi; and  carlmn   dinxidc      jlcsidcs  this  iili\  innsly  d'' 

structi\-e  iirocess,  Iiacfcria  alsu  |icrl'oiiM  a  nscinl  I'uiictiiin  in  tlic  diuc^tinn 
nl' carlioli>di'ates.  in  tliat  certain  strains  nl'  tlirni  are  alile  tn  ihuest  cellu- 
lose, I'oi-  wliidi  no  special  eiizyiMc  is  pmxidefl.     I'laelcrial  di'ie^iinn  is  enn- 

sc(|ucntl\-  essential   ill   lierlii\  oi-niis  animals:  it   takes  place  in   tin iim. 

uliicli  is  enoniiMiislv   de\e|ni.ed  I'or  thi-  piii|Hise     pai;e  -lli:;   . 

Absorption.  Tlie  ._ducose  and  levuluse  pnuluccd  li\  di'-;estinn  are 
alisnrlied  into  llie  Mi.ud  ol'  tlie  poilal  s\s1em.  Wlieii  a  very  lai'ee  (|uan- 
tily  of  a  disaccliaride.  sueli  as  cane  siiirar.  is  present  iii  tlie  I'.mmI.  a  certain 
aiiiiiiint  ot'  tlie  sUi;ar  is  alisoilied  unelianu'ed     tli.at  is  |o  sa.\ .  as  cane  suL'ar 

and  aiipears  in  tlie  liloml.  fmm  wliicli.  since  it  is  an  alnmrmal  coii- 
slitiient.  it  is  cNcreted  uncliaiiiied  in  tlie  urine.  Tliis  alimentary-  i;l.\  co- 
siiria  is  particularly  e\  ident  wlieii  llie  sin;ar  is  taken  witlmnt  any  otiier 
t'lidd:  tlius.  al'ter  takinir  cane  su«_';ir  in  an  auioiiiit  corresp.iiidinLr  to  .1 
'-'rams  jier  Uiloirram  liod.v  wciudit.   i'     \as   fnund   in  one  and  a  liall'  hours 

al'terwaril  that  the  urii t'  ten  out  .T  seventeen  healthy  individuals  eoii- 

tiiiued  cane  siiirar.  The  urine  of  three  nf  these  men.  however,  alsn  con- 
tained invert  siiu'ar  that  is.  dextrose  and  levulose.  Cane  sui.'ar  con- 
tinued  to   he  excreted    for   from   six    to  seven   hours. 

The  Sugfar  Level  in  the  Blood.  While  no  ahsdi  ptimi  of  snuar  is  o.,iiie- 
on,  the  percelitaire  nf  this  slllistaiiee  in  the  liliHid  of  the  portal  vein  is  the 
same  as  that  in  the  svstemic  circulation.  iMirini.'  alisnriition  the  former 
hcciniies  perceptilily  raised     to  what  extent   we  can   not   say     and  in  the 

latter  a  less  marked  increase  id'  siiy;ar  em ntralinii  is  usually  detect.iMe. 

Hvideiitly.  then,  hetween  the  iioint  at  which  the  sin/ar  is  alisurlied  ,ind 
the  lilooil  of  the  svstemic  circulation,  sniiie  harrier  exists  which  huld- 
hack  siinii-  nf  the  excess  of  ahsnrlied  si|e;ir.  \Ve  ha\e  very  inai'curate 
information  as  to  liow  eflicicntlv  these  harriers  hold  hack  the  excess  of 
alisorlieil   glucose  lieeause  ol    the   technical   diflieulty    in   collectiiej-   Itl I 


ilij 


if -I  '■' 


■  ifu  n 


I* 


i\.  (.-, 


'  pi   ' : 


Ir^ 


ill 


(;:)S 


MITMliirisM 

fnnnti,,.  ,,.,r,nlNnn  uitl,„„|  s.Miu,,.  .listurl,,,,,,...  „.  ,|„.  ,„i„,. 


tlil'     o 


ll.        llullM'll. 


'--  -ly  ,w,,   I.,  .WuH.   ,1,..   ,,n,Mnn   l.s  l,....n   a......,Hv   s,„,,in,   i,  ,, 

': T'r''\ '  -'^  •'■'•»-••''•  '•i-.l..i..n.i.lMl.t-.,ni,..,s:,i 

'•''■7 '■''■;'''  ''':'''^' ''''•"'•!"■'••-''  VI' IM. ■  ,|..x„.„s..  in, .„•„,., 

>iM,ill,T  Iiimim-Ihs  of  III,.  i.,,rt-il   v,M„    I      I,,     ,.   1 
.  '""■"   ^'"1         III  Midi  ..s|M.r„n,.,,|s  It   hiis  iHvn 

"'■'.""     ""■>  —  '■■'-"•■    Ml.,.,,,    i.    ,.     ,i„|,.     |_     i„     „,„     ,,, ,.„.„,„ 

'"•■■'-"•^"-"i"-,,,,,,.  „..,,„  ,„„„„„,„ ,,^,„. 


■  ii-' 


■■c     •..,».  1-.-    ,f    ,JtUf.cs,   5.,„r 


-w.,     J-    ^    'y  Jo  ts  io  Si  *c  v^ 

I'li;      I—       Inrv.  .     .l,..„,,u.    ||„. 
'~     -.t     ,,111     ,,,1^1, 


J"    -5^    «0     tJ-    «     /J     10    f 


J  fi  ri- 100 


•^'"    '""  .    '.    I'l.i.  .    '[..It., I    |i|,,.,  inn,, I'M-     In,.     /./'.,     j,,-jn- 

„,;;";;"''•'":■''"; •;•""•  "'■"'""■'>•'"■-"■-•"  .1 ..,....,,.,1,^  is 

u-JuJ:  :jr::z:::^^^^^^^^^^^  in...... 


■i"'-<  "i;il    .-ii.c  i„,t    iiM.lcr  ti„. 


Il,(,lll'l,('<'    llf    illlos- 


HS 


i'i} .. 


k''  ■,\ 


Tin:    Ml  TAI'.iil.rsM    (iK    llli;   rAHInillYI>K\Ti:: 


(;r)0 


llii'si.'i.  Sui'li  olisci-Mitiiiiis  1j!1\i'  liciii  iiiiiili'  cm  niliMts,  iiiiil  ii  I'fw  on  iiiiin 
liiiiiscir.  l*y  i-iillccti:in  lilodd  I'niiii  llic  rm  \i'iiis  nf  iiililiits.  it  lias  Im'cii 
t'diiinl  tliat,  alter  uiviiiy:  liuin   iwci  In  ten  mains  nt'  tflncdM'  tiy  stuinac-li. 

llic   <;llH'i(SC   (•iiliceliliatinll    III'   llic   sysliln'n'    IiIhikI    liririllS   to   iIm'    ill   fil't('(-ll 

iiiiiiiitfs.  attaiiiiii'-'  a  iiiaxinniiii  in  aliuiit  an  liiMic  and  tlii'ii  irtiiiiiin;;  tn 
till'  iioi'iiial  li'M'l   ill  nliciiit  tliri'c  Imhu's. 

Similar  results  liavc  Keen  (ihtaiiieil  hy  esiiiiiiiiatinii  ul'  tlio  vninns  Iiloml 
ill  man  At'ler  uivini.'  1""  ■plains  cit"  '''iieiis..  Iiv  nidiilli,  for  exaniplc.  tliorc 
is  eciinmiiiily  an  ineicaM'  in  MimkI  sinjar  aiiiniiiitiiiL'  to  rnmi  :'.0  Id  :14  l>ei- 
cent  of  tlie  normal  iiimI  la>tiiiu  for  fiem  one  to  four  lioiirs.  Tlie  existeiico 
of  tliis  piisli>niiiil'hi!  Imiii  r<ihi<  I  mid.  as  we  may  call  it.  imlieates  tliat  tlio 
suirar  retainiiiir  powers  of  tlie  liver  and  niiiseles  are  not  siillieiently  di'- 
Nclopi'd  to  iirevent  tin-  aeeiiiniilation  of  some  of  tlie  alisorl>ed  siiirar  in  tlie 

-vsteiiiic  til 1.     Wlii'iievcr  tills  increase  eseeeds  a  (M'ltaiii  limit,  some  of 

tlio  snirar  lieijiiis  to  escajie  tliroiiudi  the  kidney  into  tlie  urine.  ]iroiIiieinir 
L'lveosiiria  fiiint iiyiniiliii}  iihirii-nirid.  Tlie  pereentaL'e  of  Mood  siiirai'  aliove 
uliicli  irlyeosnria  oei'iirs  is.  in  the  ea^e  of  man.  jirohalily  ahont  0.10  to  0.11 
L'lii.  per  cent,  .\fler  daiiia'_'c  to  the  kidney,  as  in  nephritis,  or  in  loni:  staiid- 
iiiL'  eases  of  mild  dialietes.  the'  peri'enta;.'e  may  prohahly  rise  eoiisiderahl.v 
hicher  in  ihi'  liloocl  •\itlioiit   evidelii'e  of  udycosuria. 

Value  of  Blood  Examination  in  Diagnosis  of  Diabetes.  The  <letermina 
tioii  of  tl  e  amount  of  iie_'estc<l  earhohydrate  ie;|iiircd  to  liriii<r  ahmit  jiost- 
prandial  j.d.\  cosiiria  constitutes,  as  we  have  already  seen,  the  so-called 
assimilation  limit  for  sii^-ar.  which  is  often  taken  as  an  iiide\  of  the  siiirar- 
metalioli/iinr  jiower  of  the  organism.  It  is  evidint.  however,  that  the  time 
of  onset,  and  the  extent  an  I  duration  of  postprandial  liy))er!;lyeemia  must 
scr\e  as  ji  mole  certain  iieiex  of  the  siiirar-retainiinr  i>ow(>r  of  the  livei- 
and  mils'  >;  and  now  that  a  simple  and  rapid  clinical  method  exists 
'  I.ewis-Hciiedict  method ~i  for  the  acciiiate  delermination  of  sinrar  in  small 

(jllantities  of  Mood,  there  is  no  li'asoli  wli>-  this  index  should  not  he  usecl 
for  the  deteclioii   of   failiir.'  liowers  to   melaholi/e   earliohydrate. 

Ill  no  disease.  )iroliahly  not  e\  en  in  tiiheiciilosis,  is  it  more  important 
than  in  diaheti's  that  an  »'.-irl,\  diay:iiosis  should  he  made.  Thus,  if  we  find 
that  the  jiost  piaiidi.il  li\  pci  ;:lycemia  after  a  certain  aiiioiint  id'  earlio- 
h.\ilrate  develops  to  an  iiniisually  lii'_di  decree  and  persists  t'or  an  unusual 
lei|._'tli  of  time,  we  ale  Jiistilied  in  ciiitailinii:  the  earliohydrate  supply  so  as 
to  hold  thesi'  values  dow  n  to  their  level  in  inn  mal  individuals.  It  is  almost 
certain  that  the  fiist  siiiii  of  dialietes  is  an  unusual  deu'ree  and  duration 
of  jxisl  prandial  hyperiilyccmia.  .\t  liist  the  excess  of  suirar  leads  to  no 
,!:..,..,._,,.  .,,.,1  It  is  insi'llirie!!!  to  cause  an^'  eviileni  irlycosiiria,  althoiiirh  it  is 
(plite  likely  that  if  the  urine  in  sucli  iinlividuals  were  colleeted  at  very 
fre(|iieiit   inlerxals  after  catiiiv'  c;iilioli\  drale-iich   f I.  yliicose  would  he 


ill 


i 

PtI 

If 

'1 

1 ' 

■'  i 

!^  ■)' 

!    .1 


I  I  Hi' 


I       I 


fl 

(ir 


660 


MI'TAIKir.ISM 


;^"'  "'"   ;•';"•''' "■"^"■•— ^-  -'"til  tlH.  p..s„.,. i,.|  |,v,H.,.«lvn.n.i. 

:';;",'■""""""■  "";■■"•"  •-"•• iMi...  ,:;„,.;; ,r::! . 

':'V l"'nnnn..MtI.v  nn    -l.  s„  ,1.„   i„  ,,,„.„,  ,„„,,,..,,  ,„ 

liiiflivr  ixm.MtM-..  of  sM.,.,-,r  is  fciii-l. 

The  Relationship  Between  the  Sugar  Concentration  of  the  Blood  and 
h    Occurrence  of  Glycosuria.-    Ch.,..,..  ,,,„. .,  „,„  „,,,„„,  ..„,  ^^ 

'  ""'■'"'""■  "|.I....r,.M,...  .„•  any  „f  ,1...  .,.,.„•  in  ,!,..  „,in...  or  .t  l-nst 

-  1  „„r  ,.  ...■..•I,..,I.  as  ...  l.n..  .....,,  „,.,  s„,,,.,.  „l,i..h  ,.„p,,,,s  is  no,  ,,11  of 

'I'    '•v.-s  l,u,  only  „  sn,,..ll  part  of  i,.     This  ovHlow  l,vpo,l,..sis,  as  i,  is 

;:,'■;'• '"" ";". '"'"  ."•--'■-iiy  —pi.-i  i ...so  or  ,1,.  „,anv  ,vs,.i,s 

;,    "•     ■•'  '■  '""■:'  <-;""nm'.v  ui,l.  i,.     Many  of  .In...  .-x.-ptional  ,vs„I,s 

,      '  ' "  '■'"".""■•'  -'^  •'"•■  •"  "It.'-'ations  in  ,1,.  p..n....al,ili, v  of  tho  ki,ln..v 

-   -....;.  ..,.,„  ,,.„,,,,  i,  i,  ,„,,,„,.,,  ^,.,,.  ,_,  ,,|^;^^^|^_  ; 

li\  )■■■•!, ..SIS  uith  ....itain  ifs..ivati..ns. 

""•;/-"  -"■•1<  "I  \V.-.Iva.,  an.l  his  ....llal.o,.a,o,.s.  wh..  ha;'  sLnvn  l.v 
'■;""" "^   ""'•—....US   .1U..OM.   in.i....,ions  ,hat   as   ,n„..h   as  .18   ^nn    o> 

'""--""'"•   -..V  .ly,..,s,.,.ia.  al.hon.h  nn,!..,.  sn.-h  -on.li.ions  a  vc' 

'''^"""'  ""■'••■"^ ■'•'"-  i"  '""•  l'.'.v...„a^^>  .,f  s„..n.  in  tho  I.I0...I 

''';■-"'-"  th..  fail,,....  of  .l......s.>  to  P.SS  into  ,ho  ...in.,  n,,.]..,.  ...,,„,, 

'"""tn.ns.  „  has  I,,...,,  sup,.os,.,|  l.y  ,s.>v..ral  i.,v..s,i,.„.,,s  ,l,a,  ,h..  .^hu-os.. 

ppts  ,„  so,,,.,  lona  of  .•l,...ni,.al  ...,u,l.i..ati ,|,..  Moo.l.    This  ...nnp..n.,.l 

s  M,..v..    ,0  ,,..  av..  liU.,  a  ..., ,.    o f  ,1,..  , .t  snppo,.,..,.s -If  ,his 

,":  '•  "'';:  ''-."'--..,1  ,ha,.  .h..n  .h...os..  is  in.i....,...l  intrav,- 

"    isl.  .  .t  .•.I..S..S  .l,n,-,.s,s  as  U..1I  as  .ly..os„na:  wh,.,...as  .In.-os..  inj,...,...! 

: '"•"':"":'-^  "■• ""'"  '•■ '""""'  "'•'"■^"^  "^''"""-  "*• ""-  .-."-li.i  ..s  t 

;;:'::""• ''"^ '■'"'"::' \"" "•  — ^~.Mi,....af,..,.,h..a.l...inis„.ation 

I  th..  sn.a,.  a  -hs,,,,..,  anu,-ia.     To  .splain  ,h..s..  ,Ii,r..,..n....s  in  iK.havio,- 

:'""""  '■''""."T  "'•'' •^'••'•'■''  i"t.av...,o„sly  an.l  ,l,a,  ,ak..n  i„  .„i,.,.  ,,,vs 

.^s,.pp      ,,  ,  1,  ,h,..ln..os..,ao,.,..„,,i,.  p.ssin.,h.......h  the  in,...-...    .^ 

!'"■''   '"    '"""^  •'^'"'••''''•'   '"'•  ""•"'■|H..ntion   into  an.l   uti!ixati..„   L   ,(,. 

.ssn.s.  ..in..os..  ,n  a  Ur.  s.at..  lM.in«  in.-apai.l,.  of  utili.ati.,,,.     This'.-o.n- 

'""""'  "  "'"" '  '"  ' '■  "  -l'"i''"l  ".-".Mv.  a.,,1  ,h.  s,.l,s,a.....  uhi..h 


H\ 


Till.    MITAIIOI.ISM    OK    THi:    (MIllollVIHtATIS 


(iCl 


I'liiiiliiiii's  with  i;lii«-i)M'  to  I'm  III  it  i>  licliix  cil  tn  lie  rcliiti'il  t<i  tlic  'iil<'l'ii.'i1 
si'iictiiiii  III'  tile  |):ilM'i'i':is  (  m'i'  |iii;;('  t)(ii>. 

Tlio  (lirticiiltN  ill  ('xpliiiiiiiii;  wliv  tlif  lihicnM-  i,(  tlif  MimhI  ilnrs  nut  rmi- 
Nliiiitly  leak  iiitii  till'  kicliicy  i>,  liow  cv  rr,  thr  i  >\\y  ex  IcIi'ikt  ii|ic>ii  wliii-li  tin' 
li.\  |iiitlicsis   III'   a    liliMiil    Mlsjiir  ('ii|ii|iii|lllil    ri>.ts.      Nil    iliiMiiii'al    ex  iilriicf    ran 

lie  iifTiTcd  in  sii|)|)(irt  ot'  Midi  a  \  ii'w  ,     On  tl iitialy.  ail  CNpciimi'iital 

\\  ink  inilicatrs  tliat  tin'  ^ii!.'ar  (■\i^t^  in  a  I'lcr  statr;  I  nit  iinrintiinali'ly  i'mii 
this  f\  iili'inT  is  tint  riniv  inriii'.'.  Thus,  it  has  lici'n  run  in  I  thai,  w  hen  spci-i 
ini'iis  lit'  pcit'rctly  I'rrsli  liliiml  arc  plan'ij  in  a  scries  iil"  (iial.\/cr  sacs  mis- 
pcinlcil  ill  isiitiinic  saline  suhitinns.  cadi  suliitinii  cnnlainini;  a  sliu'litly  ijil'- 
t'cniit  percentage  nl'  irliicnsc,  dilTiisiun  nl'  iiliicusc,  in  one  ur  other  ilireetion. 
ncciirs  ill  all  III'  them  save  one  iiaiin'ly.  that  in  which  the  pcrcentap'  ol' 
;.'lneosc  ill  the  llniil  oiitsiile  the  dialy/cr  is  exactly  cipial  to  the  total  siitrar 
content  ol'  the  hlooil.  Such  a  rcsnlt  can  he  i'\plaincil  only  liy  assniiiin;;  that 
all  111'  the  snirar  in  the  lilonil  exists  in  a  I'lecly  ililViisihIc  state  In  its  ircneral 
Mat  lire  this  e.xpeiiincnt  is  analo'.'oMs  to  that  liy  w  hich  the  I  elision  or  pal'tial 
Iircssiire  of  to,  is  ilctciniincil  in  lihiod   (see  pa^re  '^iSi. 

It  has  hecn  assiiiiicil  hy  many  clinicians  that  u:l,\  I'osiiria  ma\   Miiiiitimes 

hi me  ijevelopcd  lieeaiise  the  kidney  fails  to  hold  hack  the  liiood  siii^'ar 

cM'ti  when  the  pcrccniam'  is  not  ahovc  the  normal  soeallcd  n  mil  ilia- 
liilis.  For  ihc  diatrnosis  ol"  this  condition  a  comparison  must  he  made  lie- 
twcen  the  sUi.'ar  concentration  of  the  lilooij  and  that  of  the  urine.  In  order 
to  do  this  at  least  two  s;iniples  of  IiIoihI  must  he  taken,  one  of  them  at  the 
heij:iiiniii;,'  and  the  oilier  at  the  end  of  a  period  diirini.'  which  iiriiic  is  lieiii;; 
collected.  .Merd.\  to  lind  that  one  .s.imple  of  li.lood  eollccted  het'orc  or  after 
or  diiriiiir  the  iieriod  of  urine  colleetion  contains  a  iiornnil  |)erccntay:i'  of 
simar.  does  not  iicecssarily  indicate  that  ;it  some  other  period  while  the 
mine  was  heiii";  produced  a  tcmporarx  liypciirlycemia  may  not  have  ex- 
isted. 


rf 

•  t 

It 

■  r 

1 

■ 

ii' 

1:^ 

< 

f 

'H 

' 

ill 


1    ii 

;•        B 

!.         fil 

'     !'■ 

'      '                i 

-n 


m 


I 


t    i 


i  I 


:  : 


I  i 


i!i' 


ni.\iTi;i{  i.wv 

TIIK  MKTAItoMs.M  n|'  Till:  <AI{|tu||V|,|{ATi:s  iC.nf.Ji 
PATE  OP  ABSORBED  GLUCOSE      OLUCONEOGENESIS 

„     ^\'.'     """      ",""      ^''''■•'     "'•■"     I "-    "••    tlM-    SU.,,„.     ,1,,,      U     ,,,,,i,„,,     |„ 

'"'    '';'■'■   '.""'    '""^'■''"        '•■""    "'i'"-^    •"''>     ■'■■M.I-M    to    it:       I,    ,„,.,v    I „. 

"'"'■'"'•  '"■  "  "'••'>   ' • ^i'li"'l  •■•■  s|,lil  „;.      Ul-  ||.,.s,.  ,„„,...,„.■.   stnr; 

'"■••'""  !"  '"•"'  ""•  "^"''  ""'I  -I-.  "iHMvas  ,.xi,h„i..n  ,„,  ,;,,,i„,,"i,- 

";"  •■'""■'■'>   '"  ""■  "' I'--  ••'"' ■-'"'  •• •■-.mi ..I'spll.liMir  ..r'tl,.. 

•-'Illr,,s,.   inni,..,,!..   „i;iv    ,|U .,-,|r  i,,   tl„.  jix,.,- 

Storage  of  Sugur.     I',,,-  ,1„.  ,„,...,„  u,.  .I,,,||  .,,,.,,  ,„..  pmn.ss  ,.r 

M,.n„r..  ..    sut:,.,,-  ;„,.!  .I,.,V,.  , M.l,.n„i„„  .,,•  i,,  „tili/.ti„„  until  „fur  ,,,• 

''•■'"■  '","'"•'';  ""•   ""'>    ""■   '   '""■'•  "f  '1-   |.. -  l.v   nl,v!,   tl„.  M.„,,... 

'-•-irs.  I.Mt  also  .|„.  i,„M,..,|,     ,.  .i,.s,iny  „f  ,!,..  sl,.,v.l  sM.,^„•      Tl,.-  stun,.',. 

",'  ^"f"'  ''•^'  ""•  "^'■'    i^  '"■'""-'•"  "l"'Ht   I,y  its  v-,M.,M   int„  a   ,,„lv.M^. 

.•l,;ir..I..  .-Mllr,!  ,//..-/'/,.     Art,.,.  ;,„  .MiM,.,l  |,.s  1,,.,.,,  .l,..„I,in^  1,,,-..  ,;„.„ 

t'tl-s    nf    ,    u,.„M..    ..,„    „,.i.|i,i..,|    „,,.„,     ,.,„.^,.,,    ,„.    .,    |,,,^.,.  ^^^    ^_j.    ,.^^_,    ^^    _^| 

";'""■''""'■  V"'"''' '''■■■'"'  "'"  '"■  '■'"""'  ' '"■•-'  •'"  '-"•'•  -"-'.■"■  thn,,  ll,;,t 

:'V'  ",","""'  ""'•■     ""  ""■  '■""••  '"""'    "  "ill  ' '-•v-.I  tl,..,t   II xtr..,,., 

IS  iHL'M.y  ..,.,,l,.s,.,.,„  ;„„I  V  i..|,|>  „n  ,1m.  ,.,.|.|i,i.,„  „f  ,,|..,.),.,I  ,  .,,.,,i,„„      ,„.;    . 

"",'"•  "';"•' ''"'"'"■'•  l'"H(i.-..,ti„M  ,.,.„  n.,-,.lily  l„.  .h.nvn  t-  -nMsis,  „f  •, 

|...!v>n.M.|u,n,|..     ,],.,  ,.  ,„  ,,, .  .,,•  ,  .„,„,,  ,i,,„  ,„,,^,^, ,^ ,,..,,  ^^^^  ,^^.^1^.^^^ 

VMS  u„I,  nun.TMl  ;H.i,l  l,..,..„„..s  ,.Mtir..!y v..,t..i  i,„„  snuMr.     If  ins. 1 

"t   rrnrnv,,,..-  ,|„.  liv..,-  i,M„H.,li.,..|y  .ft..,-  .Iratl,.  i,  is  ..,||.,„,.,|  ,„  .,,.,n.l  for 
-nu-   t,,,,,..   tl,..   yi..l,l   .,f   .ly...,....„    vill    .,.,.,.,t|y   ,|i,„i„i.,,    „„,    ;„    ;,,       ,.„.„ 

"  III  ^ipi-n-  l.r.,.  ,n;,n.i.i...  „r  ,Uu-u.,,  i„,li..„i,„  „,„ „,„„„.,  „.„^, 

;_x,st  „1„..|,  aM:„.ks  ,1,..  d,,.„^„„  ;„•„,.  .,,.„tl,  .,,,1  ...  „v..,.,s  i,  i,;,„  s,,^.,. 
Ill.s  .■M.y,„..  ,s  ,.,.|I..,|  „l,r.,„„„s,.  Tl:..  ..xis,,.,,,...  ,„•  ,,„sf,n„rl,,„  „U„;. 
Uou.h,.,..  ;.s  „  is  .....II...].  w.,„I.I  s,.,.,„  ,.,  in.li,.,.,,.  tl.Mt  .hnin...  litV  .U..  tlwr.. 

'V^  -'"-•""♦''■-' .V    I'-.r   tl„.   .Iy....,...„   i„   „,..   ,i,„,  ,..   ,„   ,,„.„.^,„,   ,,^. 

^'I,v...,.^..„..,s,..  I„„  ,!,,„  ,l,is  is  pr..v..„t,.,l  l.y  .•,„„|i,i.,„s  ^^  |,i..|,  .1,., |  „„  ,|„. 

V't.i   .n.,..H.y   „r  tl,..   liv.M.  ....II.      It    is,..i.i.,„t    tl,,„    if  .„v,l,i,„  sin.ul.l 

'".'I'l',""  ''"'■""-  '"■'■  '"  !"••■'•'••■'•-  ^^i'li  Iliis  i„| ,i„.  i,„|„.,,„,,  „„,.,,,, „ 

Ul      l,.>r„nH.  ,...„v..|-t...|   int..   ^.Ju.......   „l,i..|,   .,„   ,..,,,,,;,„_,.   j,,,,,   ,,~;  ,,,,;,„, 

will  p|..»liic..  Iiyp..|;rly..,.nii;,  ,,11.1  -,'l\v.  s;iii.i. 

Sourcesof  Glycogen.     I„  s,„.|y,;„.  ,|,..  s„„,.. ,■  .,„_„,  ;„  „„,  ;,„,,„,, 

'HHly  It  IS  „Mp,„.,;,„t  il,..,.,.f„,,.  ,1,,,  „,.  s|,„„l,|  (i,,.,  „,.,||  |,„„„   „^,„.j,^.  ,,,,, 


Till.    Ml  TAIlol  l-\l    iiK    nil     I   \llliii|IV|>H\ll  s 


(it;:i 


I'dliilitiiilis  llliilr?-  wliicli  ulvrMtri'ii  iiiii>  111'  fcHiiii'il  ill  till'  liver;  llifit  is, 
wlirlliiT  it  is  t'liniird  fiuiii  iiliviitli<(|  milmi  hIcimi-,  ni  \'\i>u\  ntjur  >iilislaiii'i'H 
lllsii,  slli'll   IIH  pl'iitrili   iilnl    fill.      Till'    illl|i'irtillHT   III'  Mirll    Klmw  Iiiliri'   ri'sfs 

ill  thi-  I'lii't  tliiil  ill  M'M'tr  ijiiilii  tt».  -ii'.'iir  still  I'lPhtiiiiHs  til  ln'  iiiMiil  III 

tllf  III 1,  ilillli'llU'll    III'  MIUMI    is  liiillU'   tllUi'll    with    till'    I'iMiil        'I'll  rlli'i'k    till- 

liyixT'.'l.v  I'i'llliil  ill  slli'li  fuses  it  liieiiliies  In  eessiirv,  tliep  el'me,  tii  ellltiiil 
the  iliet    lliit    nlllv     \vilh    li'lfll  111   I'l   its   I'ii  llmh  Mllille   eulililit.   Iillt    illso   with 

li'.'Ill'il   to   U  hiitiM'l-   iilher    fuiiiNtlilT   lnil\     1 ilpillile   i.t'   e.iU^iliir   ulveiiueii 

t'lirmiltiiill.  'I'lii'  |il'iietieiil  i|l|i'Ntiii||  tlielel'mi'  i»,  Whilt  lire  tlii'-e  riiiiilsllllTv  ' 
Tliere  ille  t\Ml  llli'lhnils  liy  whiih  tie  |i|iililein  IIIM,\  I''  ill  vest  hj-jiIi'iI.  'I'lie 
tilNt,  wllieli  we  111,'IV  I'iiH  the  ilireet  iiiiIIhmI.  eulisisU  ill  I  einleliii'.'  the  li\i'r 
fiei'  iif  Klveie.'eii  ;ili<|  llii'll  snllie  time  ;|  I'lerw  ill  il  I'leilili!.'  the  iihllilill  \vitll 
till'  riiiiilslllIT  ill  i|llistiiill,  iiriel Willi]  killillir  it  flllil  I'Nilllliliill'-'  the  livi'l- 
I'lir  i.'lM'iii.'eli  The  iillier,  v\liie|i  we  liiiiv  ciiil  the  iinjileel  IMelliml.  I'l'll- 
sisls  ill  (il'st  111'  iill  leliclirilii:  the  illlilMllI  illi'll|ii|lil<'  111'  Hxiili/illU'  i-'lueiise 
thllf  is.  IllilUilli,'  it  iliillietie  illnl  ihill  iniieeeilillir  In  ^i'"'  \i  In  tiler  the  ill- 
jjeNtimi  111'  il  nix  ell  I'limlstllll'  eiillses  iiii  ilieicii-e  ill  the  Mliriii'  e\e|itiii||  ill 
the  lllilie.  'I'lle  lllithnils  I'ltf  lemlerilii,'  iili  ,'lliilliiil  ev|ieiiiiieiitiill y  c|i;ilietii' 
\\ill  he  (•iiiisiileiic!  hiler;  fi,r  the  iHiseiit  it  is  iiii)iiii'tiinf  to  Mute  thill,  it' 
il  iliillietie  illlililill  exeretrs  iimmc  'jllleiise  while  feij  mi  il  '.^Mll  I'l  kmIsI  IllT. 
we  iiiii.N    infer  tliiit  the  iinriiiiil  iiuiiiiiil  wmilil  emnert  it  iiiln  i.'l\  ein.'cii. 

The  l-isiilts  iif  the  direct  inethiKl  ille  IMliell  Ic-s  leliiilile  tlnill  th'i>e  of 
the    ilnlinet     for    the    iei|v,,||    thilt     il     i-    i\t  leiiiely    ilil'tiellll     to    feiilnve    III! 

tl'iiees  of  '_'l>e.ii;eii    (v the    li\er.      The   liiethocN  ellllihi.N  eil    for  this   |iiir- 

l»os('  hii\e  eoiisixteil    ill:      i|'    st.irx  ill  ion   of  the  iiniiiiiil:   I'J'i    iiiilM'illiir  I'X- 

ereise;  i  :1  i   cNeieise  iiiiil  stiirvatin iiiliiiie.]  ;  iiinl      i      the  |)rnil'ietion  of 

eeltaili    I'olllis  of  cxiieiiiiii'litiil   iliiiheto      for  (AiilM|>le.   tlmt    iirmlii 1   hy 

jililorlii/iii.     Stiirviitioii   nlone   is   niisiitisfiietiry.    lor    it    hiis   heeii    foiiinl 

thilt,  iiltl'.oimh  iit  eertiiin  •^\:\i'   ■   "f  this  • lilioll  the  li\er  \nn\    I me  ill 

most  entirely  free  fron,  iiny  n'.iee  of  •.rlyeoueii.  ill  il  liiter  stiiiii-  u!.\ein.'eii 
mny  iiiriiin  iinilM'  its  ii|i|ieiiriiin'e.  It  is  therefore  most  iliflieiilt  to  ileeiile 
iit  whilt  s1;i<:e  in  stiirviition  the  iiliilllill  slloiiM  '■»■  eoiisiilereil  iis  l'I.v  eoiieli- 
free. 

If  the  stiirvini:  iinimiil  is  iiiinle  to  )ieil'o' in  ;  niseiihir  exereise.  eomiiletc 
reliloVill  of  '.'lyeoL'ell  from  the  liver  cilM  lie  (lepeinleil  npoii.  The  exereise 
limy  he  iiiinliiee,l  hy  the  inlmiiiisti'iit imi  of  st ryeliniiie  in  siieii  ilosiiu'e  lis 

just  to  i>roilue( iiMiKions  of  the  volnniiirv    mii^eles  wilhoiit  pei  nmnent 

eolitnietioii  of  those  of  res|iiriitioii.  The  most  useful  methoil.  however, 
coiisisls  ill  stiirviiiL'  the  iiniinal  for  ii  few  ihivs  iind  then  iiliieiiii:  it  in  a 
colli,  (liimi)  riioiii.  Jit'ter  'A'wiuu:  il  ii  eohj  hiilh.  The  eviiporiitioii  of  mois- 
ture from  the  surlaee  so  eools  the  iioily  ■low  ii  ihiil  llie  eiyeoyen  .stole  ail 
hcconies  used  up   in   the  iit1ciiil>t   to   supply    fuel   for   the   proiliietiim   of 


llhn 


!      : 


■il 


iir- 


iL:!' 


!i  :i!i  ,1 


■  1 

.'^ 

'        i 

■!, 

,i|'      i 

I               ' 

i 

\] 

ii\i^ 

lit;  I 


MITAIKIIISM 


siiriiririit    lira;    1,1   iii.'i  i  1 1 1  a  j  II   til,.   |„„|\    tciii|>,'i'atiiir.      Tin's   nii'tliiMl  ,mii   !„■ 

'■'■'"''■'■"■'I   -'ill    i""i '■'■■liii    ill   .■tr,.,-liii,_r  a    ,viii,,\al    ,,)•  all    rarli,,li\  ,|i-iit-' 

tV"iii  III,'  l,",|\  liv  -niiii..  til.,  animal  plil,,ilii/.iii  ,.\,.iy  citilit  li,,urs.  This 
ilni'j-.  as  wr  ■-liall  m-.  rciHJtMs  the  animal  dialifti.'. 

Ai'li^i-  iviii.n  iii.j   il,,.  L;l\,',i-,.n,   rnrtlK'i-  (icpositi,!!!   in   the   liver  can   Iif 
iv,i,lil\   v||,,vMi   I,,  ,1,',-ur  whrtt  anv    ,,|'  )li,.  ,n'<!inar\    Miuars  ,<v  starclics  ar.' 

"'^''"  ■■'-   ' I       ''   'I'"-   II"'   "M-nr.   Iiiiwcvcr.  \\li,.n   <-ln'mical   siilistanccs 

fl,,s,l\     ivl,-,t,',|    I ,linai-y   Miyar,   Mi,-h   ;is   tl,,.    wimil   siiumis   (liciit.iscs^ 

'"■  '!"■  .-il''"!!"!-  •■iii'l  a,M,i' i-ii's|.,)ii,liiiu'  t,,  ,1<'\I  nis...  art nlaim-il  in  the 

,li,i.  \,ir  i\,.r~.  it  ,„-.Mir  Willi  ,-cllnl,,s,'  ,,r  \viili  iniilin.  a  iHilysai-cliari.l.' 
iMiili  ii|>  iViim  p,'ht,.s.'  simar.  Wli.'ii  pnilcins  arc  \\;\  tlic  results  arc  imf 
s"   ,|,'liiiit,>.   allliiiii'.'li    many  ,,li>,.r\  ,.|s   have   cla'iii,',!    tliat     u'ly,-,,;:,.,,     is 

I'niiii,',!.      With    fat.   cm  til,'  i.tlicr  liaicl.   im   ul\i-,c.r,.||    i',,nMati( an    he 

-1'"^'"  ' '•Ill'-  :iltli"niili  u,'  kiHiu   that  a  tra,c  ,,1'  carli,iliy, Irate  miisf  lie 

r,,nii,,|  ,1111   <>\   111,.  ■.H,\,.eriiit    ,,t'  ili,.  fat   m,ilei-nl,>. 

'I'll,'  r,'^nlt^  ,,t'  tile  ilirei-t   m,'ili,„|,  ,.vi'ii   w  li,-n   tli,'  , Iiti,.ns  are  jter- 

t',-,-il\  (■.,nti,,||i',|.  ar,.  ver,\  nnielialile.  lie,-ansc  any  ii,.\v  sm^ar  in-iHlni'cil 
l,y  111,.  iii.j,>t>',|  sul,st,in,-e  iiisti.a,l  ,,1'  liein;:  >t,,r,.,i  as  -;lyeiii,|.||  mav  !),■ 
'liivelly  II-,,, 1  1,\    th,.  tissues  as  it   is  t',Miin-,|.      Where  ,inly  a  sli-hl  .le-ilee 

"''  -!ii'"i uii.esis  is  -lurin-.  it   is  n,,t   likely   th;il    any   i.T  the  ^'lue.is,. 

\'ill   Ire   retaini-,|   in   th,-   li,iily  as  ul  \  cmjch, 

Tl„-  iii,tli,„ls  eiii|,l,,,\,-,|  f,,r  |ii-,„|ii,-iiiM  ,.N|i,.|-im,.|ital  ,lialM-t,.s  In  investi 
uati..ii  ,,itli,-,.  |M-,.M,.|ii>  liy  111,,  indirect  method  a  IV  i  I  ■  the  eniir,.  r.-im.val 

"''   'II''    p.iii'-r,  as.    and    i  •_'      the    |.,iiiti iis    adtiiinist  rat idii    ,,!'    th,-    drin.' 

I'li'"ilii-''ii  'I'll''  ■■iniiiial  ren,l,.r,.i!  ilialnti,-  li.\  i-iiher  ,,|'  these  iin'tlnids  is 
first  ,,1-  all  ,.l.s,-i-v,.d  f,.r  s,-\eral  ,Iays  t,i  di-lermin,-  the  n.Miiial  daily  (.\- 
eivti,.ii  ,,|'  sin^^ar.  At  Ih,-  sam,.  time  th,.  iiitr,e_'i-n  |.\er,.tii,n  I'm-  the  d;i\' 
is  d,l,.riiiiiii-,l.  the  rati,,  l„-tueen  the  l,,lal  nitr,ii;,-n  ami  the  ■ilncisi'  hi'lm^' 
KiM.uii  as  i;   I,,  \   rati,,,  an, I  l„-iii._.  al„,nl    1    I,,  :;.i;:.  \vli,.n  ,-,mi|,|etc  .lialietes 

'i.'i-   '"'''"I stahlish,  ,1.      Th,.    f,.,„|stnrr   in    ,|U,.sti,,ii    is   tli..n    fed   1,i   tin' 

■iiiii".-i'.   •■iMil    ill,-   am, ,111,1    ,,r  I'xti-a    ■.;liii.,,s,.   e\m,.I,-,|    tli,rel,y   is   taken    to 

r.-l,r,  s,-iit    that    \\hi,-|i    has   l.eeii   ,|i-ii\,-,|    fr,,!,,   the   ii|,j,.sle,l    f |,      ]}y   fids 

iii'''l'"l    it    li^s   |.,.,  II   |„,ssil,|,.   t,,  s|i,,u    thai,   II,, I   ,,iil.\    111,.  al",v,.   m,'nti,iiM.,l 

,-;irl",li,\, halts,    I, lit    |,r,,t,-iii    as   \\,-l|    i,r,„| a    \,-ry    e,,iisi,l,-i-al,l,.   ,|nan- 

tit.\-  ,,1  'jlm-i.M-  in  th,-  animal  l!,„ly.  Fats.  Ii,,\\ ,-., ,.r.  .\  i,'!,!  ,,nly  n,'ir;itive 
r,'siilts 

'I'l'"  iii'liK'''!  m-.th,„l  has  aii,,tli,-r  •jr,.at  a,|\  aiita-jc-  ,,\(-r  tlie  ilirect  in 
'II'''  'II''  '■, --nits  at,-  iiin,-h  ii|,,i-e  iiiiiiiilituliii  in  eliara,'(er ;  I'm-  e\aiii|)lc. 
'-'i-l-  'Il '!  li-.  |iii|,ils  ha\,'  h,-,ii  al,li.  t,i  .h'termin,.  th,-  am,, mil  ,it'  yluc'is,. 

'"^''i'l'  •■III  i"'  l'i"'lii t   i>>    lee,lin..;  c-,-rtaiii  ,,t'  tin-  l,nii,iini;  sl,,iies  nf  the 

pr,,t,in    m,,|,-,-ul,-.      Th,.    -r,.at    |,ra,-tii.al    iiiip,,rtaii,-e    ,,t'   sui.ji    r.-sults    in 


Till.    MITAI'.ul.ISM    iiK    llli;   i  Al;r.(illYI>II.\ri.: 


Odi 


till"  tlirr'iipy  of  (li;ilicti"<  iiiiiki's  it  ;iil\  UaMi'  I'm'  iis  to  'jd  into  tlic  MilijiM-t 
ji  little  more  in  (Ictnil  Ihtc. 

l)ij'js  iiic  rriidcicil  ili;iliclic  li\  plilni  lii/in  t'ollowcd  liy  n  colil  liatli 
Miiil  e\|ioMii'i'  ill  a  colli  room.  Wlicii  all  ol'  iln'  oriiriiial  trl^  lo'.n'ii  in  the 
lioily  lias  licfii  mit  liij  of.  as  cv  iili  ni'c(l  liy  tlic  roiisiiini-y  ot'  tlic  (I  to  \ 
ratio  ill  till'  daily  i|uaiit it ifs  o|'  niiiir  rM-rrtcd.  tin'  siil|.,iani'«'  under  iii- 
\  I'stiiralioii  i-.  fed.  II'  tliis  Milislaiiei-  rniitaiiis  no  nitroL'cii  and  caiiM's  no 
clian;.'!'  in  tin'  iiitt'o^'cii  I'vcrel  ion.  aii\  iin-rrasi'  in  tliat  of  ylucosc  must 
(ijnioiisly  ri'|iri'^('iit  tie  extent  to  \\liie|i  tlii^  sulistanee  lias  lieeome  ('(iii- 
\eitecl  into  tliis  siii;ar.  On  tin'  otiii'r  hand,  if  'lie  siilistanee  itselt"  eoii- 
taiiis  nit  reiicn.  or  il'  it  -aiiM-s  a  elianL;e  in  tlie  i\i-retion  id'  nitroy;en.  it 
lieeoMies  neeessary  to  ealelllale  liow  liiileli  o|'  llie  e\rreted  i;hieose  mi'.dit 
lia\e  lieeii  deri\ed  Ifom  |  jic  lioiiy  |)riitciii.  as-,ni;iiiiL;'  tlial  tliis  eaii  t'oi-ni 
i;lileiise.  and   liow    mileli    I'ldiii   the  admiiiistereij   -  nli^taliee.* 

l'"rom  tlie  result-  o|'  this  metliod  it  has  Ircrii  an  easy  matter  to  show 
that  the  rollow  inu  siilistaiiei  s  are  i-nh\eiteil  in  the  animal  lioijy  into 
Ldileiise:  (  I  i  dhiiitl  iililihnd'  <'||  "'11  •'li(>i.  I'.y  plaeiiitr  three  mol- 
ecules ot'  this  sulistanee  toui't  her,  a  hexos,.  nioieeiile  will  'lie  ]>rodiieed, 
a  synthesis  wliieli  eaii  lie  aeeoiiiiilisheil  in  the  eliiinieal  lalioratory. 
(d.M'ol  aldehyde   iiia.v    lie   I'ormed   in    noriiial    metaliolisin   out    ot'  fjlycncoll 

.(irAll.rooiiv 

iL'i  Clunr.,!  (CM  fill  (  llu||  Cllnli  may  also  rea-lily  he  eoii- 
\erled  into  hexose  ill  the  lahoralory.  the  possilile  intermediary  jiroiliiets 
heiiiu'  dioxyaeeton.'  1  CM, (III  C<  I  ell  <  III  and  ulyeerie  aldehyde 
iClLtlll  CIKlll  CllOi.  Two  moleeiiles  n\  .-ilher  ol'  these  may  lie 
pnlymeri/ed  to  I'.irm  a  hexosr  Mioleeiile.  and  when  this  proeess  neeiirs 
ill  the  animal  hody,  the  hexose  ii.rmed  is  i;lue.ivr. 

(oi  l.tiriir  ,1,1,1  iCIl  ClKlM  ((Kill  I  is  complelely  ixerted  to  dex- 
trose ill  the  dialielie  animal,  and  the  pi-oiTss  must  iir.ohe  Imtii  a  re- 
arrailLiement  ol'  tlie  liioleeulr  and  siiliseipiiMit  pol\  !ileli/,it  iiii..  The  I'elati'd 
sulistanee.  priip\  I  aleolml  (11  (jj  (II  (til  is  aKo  eonverled  into 
ullleiise  in  the  |ili  liirh  i/iii  i/ed  d'U.    .\s  to  the  rxai-t   natnie  oi'  the  ehemieal 

ehailV'es     whieh     oeelir     as     intermedial  \      slrps      iu      the     louMision     of     these 

sulislaiiers  into  ulneose.  wr  iiic   hot    as   \,\    r,  liaiii.  hut    a   step  has  I n 

made  in  the  disnisrry  that  a  suhstam ailed  met  h>  lulv  oxal   '  (  '11  (  '(  M  '||(  »  ' 

may  lie  olitaiin-d    I'mm   hn-lie  aeid   and   aNo    I'rom    ltI se.  and    tiiat    this 

sulistanee  is  eon\erled  into  uliieosr  when  it  is  administered  to  phlorlii- 
zilii/ed  diij-'s.        W'e    shall   tind    later    an    iiniuMtant     role   for  this  siihst; e 

■Till-   .  il,  iil,,l ,■   ,  !,       -    1    !'..x>.:       ir,     ,,".    ■;.  .    ,,•     ,  ,,    .-   ,,    ,;.    .;..    ,-,  I,.  ,,,.   ■,  .,    1    ,-,!,., , 

'ii'iiuUmi    frc.in    tllf    -iitr..;  •  n    (\.r.  tmn      .nil    -Iw    JitT,  t.u,  ..    ut...l.    .mm-1     ,,.,r.«,.I!t    tl'.v    •; !  t  •  "i:f!:    -A    \\\:- 

Iv     ploUlPl.     I-     1111II1..I,  ■!     1.1      II.      I.     ...      \      ■.,...,     si'k,  I,  ';,,>..,!,, I     ,,-,     'lli'.'.l.ir    ]..,m''',-     |,,    "li.,"l      '.'.'•, 

"'■•"Il    llw    -nl.-l.:n,.      «,,,     l..|         W,      ,,l,l.|.;-     i!i     111.    n  .n      !■.      ■.•!i,  ....       \.-,\.    I     !,..■.,     |!,,      I.,,.iv  I',. 

.■l'irn<e  ciiilnc  fr,,m  llir  n.lmin-l.  r.  .1  .nr.l.ii:c.-  :  ni  tlmi  I  .■  i^.  crt  liiu.l  l.v  Jfliicliiiij  iImI  .liriii  I 
ni.ni  thf  hiit\    i-rMciii  tfiii  liii    i.ii.il   kIu."-.    .-.i.t...ii 


I 


il 


?  :i 


fitlC) 


MlTAIiDI.lSM 


ill  till"  I'jisc  III"  l';it  iiiftiilMi'isiii.  jiiid  it  ciiii  also  rpiidily  1)»  iiroilucod  <luriii<^ 
llic  iiitcriiiciiiiiry  iirciikilow  ii  of  ccrtiiin  nf  flic  protein  liiiildiiiii-stoiics. 
siicli  for  fXiiiii|>lf  ;is  iiliiiiiiic  (t'll  CIINIIJ '(  K  »Ih. 

TlicM'  <'li(iMic;il  |>ossi)iilitii's  rcy,ir(liii<j  the  iiatlin'  of  tlio  substances 
tliat  SI  r\  c  as  ste|)i)iin.'  stones  lictwcen  tlic  aliovo  sufiar-foriiiinir  sult- 
staiiees  aiiil  siiu'ar  itself  iiia.\  lie  translated  later  into  iirolialiilities  on 
aeeoiiiit  of  tlie  discovery  tliat  tlic  en/ynies  exist  in  various  tissues  wliieli 
are  iiivohed  in  convertiii;;  iiietliylirlyoxal  into  lactic  acid: 

fir,  rif. 

I  I 

ro  ir-^iicoir 

riio  rooii 

rin.lliUJv'^^'ll  diic-fe'   !M-i.n 


Tliese  tii/yiiies  are  eallecl  iihtii.niliixi  :i,  and  since  tlie  reactions  ■\\liicli 
tliey  UK'iliate  are  uiidonlitedly  reversilile  in  (diaractcr,  it  is  prolialile  tliat 
tlie  ciiiiversiiiii  into  sui:ar  of  lactic  aciil  and  alanine  to  take  lliosi-  two 
as  ainoiiir  tlie  eoiniiciiiesl  of  the  siiirar  |)recursors  of  tlio  animal  liody — 
oeeiirs  ai'cordiiiu'   to  the   fnllowiiiir  e(|iiation: 


fll  cilMI  rooii 


N 


l:o  HI  ■<  I'll  ('(ll'IM)  — I'.ll,  (), 

III  <ll(HHni>ll    y 

l:Hlii-  tli-l.jl  I  llirtl.vl^l\ii\:il  I     fllr\">-i'') 


-a 


'I'lie  uiiiijiie  position  of  iiictliyl'jlyoxal.  Iiesides  e\i)!ainin<;  tlie  known 
I'csoliitiniis  iif  protein  and  fat  aiol  earlicdiydrato  in  interinediary  nu'tali- 
olisiii,  is  also  of  iiiipni  lance  in  cxplainintr  the  synthetic  iirodnction  of 
".'liicose  fi'oiii  friietiise  (nr  le\iilosc'i.  I''riietose  will  first  of  all  liecoine 
<-oii\ertecl  into  iiietli\  l'^l\  o\al  radji'lcs.  and  these  will  then  lieconie  syn- 
thesized  into  u'liicose. 

The  theory  of  the  eoliMl^iiili  of  yluensc  info  lactic  acid  as  a  stciipillLT 
stone  in  the  iiietalpnli^iii  of  carliidiyiliale  meets  with  oiip  olijection: 
namely,  that  the  laetie  acid  In  not  prodiU'cd  from  earliohydrate  in  the 
or!rani->m.  e\ec'pt  in  eases  where  there  is  ox.\  ltcii  deliciene,\  or  excess  of 
alkali  in  tin'  tissue  tinids. 

Comiiii;  now  to  fhe  amiiici  aeids,  which,  it  will  he  reniemliered  repn- 
sciil  the  hnildiii','  stones  nf  the  piiilciii  moleeiih'.  it  has  liceii  found  that 
^■|\coi-oll.  ;ilaiiine.  and  aspaitie  ;ii'd  irliitamic  acids  inerea-e  the  ijlueose 
exeri'tinii  when  'iivcii  til  plilorhi/iiii/ed  iloirs.  whereas  leucine  and  tyro- 
sine ha\e  no  siirli  actimi.  I'y  the  method  dcscrilieil  aliovi'.  it  is  ])osslli|e 
to  clctcrmiiie  the  exaet  proportinii  of  the  eailiuii  of  each  of  those  amino 
acids  whieh  liieiniieN  ccoix  erti>d  to  lilncose.  This  is  shown  in  the  aecom- 
jianyin-:  talile. 


Tin:  mi.t.mi()I.i>;m  ok  tiii:  cAUi'.oiiYiiHA'rr.s 

TwiMV  (liiwis  (iK  Tin:  V.\r;iOfs  Amino  liniUKS  \Vi.i;i:  (livKX  to 
riii.iii!iii/.iN  i>i.mu;tic  Po<;.s 


6t)7 


.\VI,l;A(ii:   WllM   NT  rKlillMU.K 

OF  CI.ICIISF.   I'lIO-  (  I1AN(;I. 

IHCI  i>  IN  nnnv 

i:;.!:;    i  ii\c    .i"^s,  All  c  >  "ti\ciic.| 

iiiM'    '■■A\r    !"i.77)  t'l   i;lu('iisi' 


Cl.rcOSK  THAT 

worm  ni;  rno- 
iirii:ii  iiY  CH  WCK 


Aim  ANii  hm:mi  I, a 

(ilv .11 

(IIMI  I'OOII 

i.  ;<l;iTliii(' 

ClI/MINHroOlI 

A^|':iitii-  .M-i  I  l-.l-    i  I'miii-    th<iis)        Time   nf  llio   f"iir  i:'..."!^ 

cooll    en     ciiMi     (-(Kill  r  ;ii..Mi-^  I-  iiMit.'.i 


1- 


(  two     (1n;;s) 


20. i2 


(lliitiiinic  aciil 

cnoii 

/ 
(•II, 

I 

(•II,     ClINII 

I 
COOIf 


i:;.:;l 


Tiir  V  ..I'  til.'  dvi'  yi:i\ 

(•  riliiiiis  r.iiiMi  ti'il 
I,.   ;;lu.'i.s,. 


It  is  of  I'lirtliiT  iiitiTol  til  iHiiiit  out  lli;it  tlirsc  I'mir  aiiiinu  iiculs 
coiistitiltc  iihont  '2(1  i>iM-  cfiit  of  ;ill  tlu  amino  ariils  in  llr^li  iiiuti'in.  ami 
lliat  llic  total  virlil  of  -iliH'osi-  fimii  tln'in  conM  Ik-  "^ti.:!  u:rams:  tints 
ai'coiMitiiii.'  for  iifarly  one  lialf  of  tin-  (ill  grains  wliii-li  a  dialu'lic  aiiiiual 
in-oiJiiiM's  ffoiii   1110  L'l-niis  of  lli'^ti. 

Gluconeog-enesis  in  Normal  Animals.  Altliou'.'h  it  lia>  lucn  i-lrarly 
sliowii  l)y  tin'  imliri'i-t  iiii'tlioil  lliat  not  only  protrin  Imt  its  ili.i'oni|)osi- 
tioii  itriMJiicIs  as  well,  can  lio  loaijily  immu  iTtril  into  ului-osr.  \v\  this  iloi's 

not  iifc'i'ssaril.\-   imlit-ati-  that    a  siiuil.-n-  i-on\  iTsii.i rurs  in   tlic  noinlia- 

lictif  aiiinial.  'i'iiat  surli  is  tio  I'aso.  however,  ran  he  shown  in  various 
ways.  Thus,  at  the  cinl  of  a  i>rri.iil  of  Vwvx  starxalion  consiilorahh' 
(|Uaiititi('s  of  '.'Ivcouen  arr  (|uiti'  coiiiiM.iiil.v  fonml  in  tiir  lioil.v',  ami  the 
hlooil  snyar,  althoU'jh  lower  than  normal.  ne\er  eiitirelv  ilisapi>ears. 
\ow-,  sinee  no  earliohyilr;ite  is  lieinir  inuesleil,  ami  the  limly  stores  of  ihis 
fo.Mlstuff  he.-oiiie  cNhausteil  early  ilurini.'  slar\alii'n  n-f.  pa^e  (id:!  i.  it 
is  cviilent  that  the  eaihohvilrate  must  he  ]i|-oilr,eeil  fl-nni  the  l>rotein  of 
the  aniiiial's  lioil\.  .\  still  more  eon\  ineinir  experiiio'iit  ean  he  con- 
dueled  hy  pi'odueiiiu'  strychnine  con\ulsions  in  a  slaivin-^  animal.  If 
llie  animal   is  killed  after  the  convulsions  lia\e  lasted   for  a   ceilaiii   time. 

the     tissues     will    111      found    almost     if    mt     entirel.\-     fr if    ud.v  col'.-ii. 

hut  if  the  eonvnlsiiiiis  are  made  to  disapjiear  hy  uiviiiir  chloral  and  the 
animal  allowed  to  sh'cp  for  some  time  het'ure  killinu'  it.  elycoiv,.!!  atrain 
accuimdatcs  iii  the  Imdy.  T!;i--  j;!y. ■.,._'*>■,■,  .■:■!-<  have  heen  tii.-oiufa.-tured 
out   of  iioiicailioliydiatc  material. 

Coi-rohorativc  ovidcnco  of  a  somewhat  different  nature  is  furnished  hy 


t 


I  J* 


w 


ipi; 

ItfcJ;!. 


m 


■x 


%\ 


i 


'■.II: 


m 


1:m;:1   mi;i   ; 


i 


(;(is 


Ml   rAllflllsM 


Mil   cxjiiiiiii.'il  inii    111'   111!'    i-cs|iii-,if(ii\    i|iiiiliciit,    wliicli,    it    \\\\]    lie   I'cniiMu- 

licrril    '  |Mjrc   "liT  ' .   VMI'ii'S  : Mcliii'/   tci   till'    IKIilirc  nl'   tllf    fnuilsllllV  iir  Imdy 

('iiiiNtiliiciit  tli.'it  is  iiinlciudiiiu  iiii't.ilinliMii  Jit  tlif  time,  ln'iri";  iiliiuit  1 
\ulli  cii-IimIi^  i|i-;i!r  .-irid  .ilKitit  (1.7  willi  pruti'iii.  It'  tlii'  i|i|iiticiit  is 
iilisi'i'cil  iliiiitii:  st;ir\  .■ilimi.  it  will  urtm  lie  fdiiiid  tn  I'lill  IicIdw  (1.7.  ;i 
lii;iin'  \\liii-li  i-:iii  lie  r\|i|;iiiiiil  iimI>  hy  .•issiniiini;  lli;it  <i\.\  <ii'li  liiis  liccii 
rct.iilicil  ill  tile  \u,,\\  lirwiiid  llir  i|U:i'ili|\  uliii'li  is  ln'ccssa  !\\  I'nr  iniMIc 
ili.lli'    pllr|Hisis   (if   ii\i(|;iti(i|l    let'.    r(|U,iliii||s   im    (>:iL'r   ."i  |S  i  . 

Siii'-r  it  is  kr.iiwii  tli,-il  ill!,  n  tiiiiicil  oxyirrii  <  :in  iint  exist  in  tiie  Imily 
ill  ;i  I'l'''  sliilr  jl  iMIlsl  l.c  cnMi'hlili'rj  tililt  It  liMs  I  ii'iM  iliic  i  lliMirpnf;it  ci  I 
ilit'i  slllist.ilicrs  h.-uili"'  ,1  liiiili  ii\.\L;rii  cMiitclit.  Slleli  Would  In'  tlic  imm' 
it'  |iiiitciii  (ir  t'at.  wliicli  ci'MtMiiis  niily  t'lnm  I'J  to  'JO  per  cctit  nt'  cixy'^cn. 
w  I'lc     ciiiiN  ifird     1"    i-:iilinliyi|i;itc,     wliidl     riilit;iilis    ;ili(Hlt     ."):!     piM'    crlit. 

It  ili/;!tinii  i.i'  iiiliidid  i.w'jrii   I'm-  this  piirpiisi'.  as  wc  liavi'  si'i'ii,  1 mii's 

\riy   stiikiiiL'   in   thr  ciis.'  ,.t'   liilirfiiatiiiLT  animals  diirini;'   tin'   wintrr  sli'i'p. 


\  ''    i' 


m 


1     i 


Mil-- 1  ■!  '^^ 

1 M        I  '  f' 


CIIAITKI!  I. XXVI 

Till-:  Mi"i'Ai!<»MsM  <»r  TIN-;  (  .\i;U(»iivi)i;ATi:s   (nutM) 

FATE  OF  GLYCOGEN 

llii\in'_'  Iiccdiiit'   fiiiiiiliiir  witli  tlic  snui-ci's   rrmii   wliic-li   ulyn'.'i'ii   iii;iy 

he  cliMi\C(l.  we  iii;i_v  lltiw  pidcccd  tii  stui|\  tin'  I'.ltc  nT  tin'  trl voiri'll  fniiml 
ill  lllC  liver  cells  iiml  in  tlie  iiillseles.  Vi<v  tlie  iMi^^riit  ve  sliall  emifiiie  oui- 
Mtlelltioll  ti)  llie  !:ly"'-''ii  "''  ''i''  liver,  ir  ;.  pnitimi  of  li\ei-  reiil'ived 
tVdIil  il  '.vcll-l'eij  ;iMirii:il  is  exiiinined  iili<'l'ci-e"pie;;lly  :it'ter  slniliillL'  eitliel' 
witli  iiidiiii-  or  with  eiiniiiiie  hy  I'.est '-.  nietliciil.  it  will  l>c  round  tli:it  the 
eells  of  the  loiillles  ;ire  tilh'd  with  lilyeie^m  exeept  lor  the  lilflei.  whieh 
lire  free   from   this  sllhstjiliee.      It',  oil   ihr   .itlo'r  hlilld,   the   li\i'r  is    fr.illl   nil 

.'lllillllll   tllilt  lins   not   heell   r •ntly   fed.  the  loliules  will   eo|lt;iin   no  -jlyeo- 

e-en  esceiit  for  :iii  ;ireii  lim'ih'.'iiiu'  on  the  eeiitriil  \ciii  ;ind  i>i'rh;ips  n 
n;in-o\v  strip  iit  tlie  jieriphery  of  the  hihiile.  When  it  is  jne^ent  the  reln- 
ti\e  Jiniount  of  <.dyeoi;e||  ill  dilTeri'llt  hihuh'S.  lis  drlenilined  elieiiiie.-illy, 
is  the  siinie  ovi'i-  the  entire  li\er  tlnit  is  to  s;iy,  no  one  lolie  is  rielicr  in 
this  snlistiinee  tli;iii  jinother.  Xothinu  detiiiite  is  known  ;is  to  how  the 
u]yeo<;en  is  lield  in  tiie  |n'otopi;isiii  of  the  c'lls.  ;ilthoii'_di  some  histolo- 
u'isfs  snu'irest  tiiat  it  is  eonihiiieil  with  Ji  siisteiilaeiilar  material  especially 
pro\i<lecl   fen-  tliis  imrpose. 

Tiie  >jl\-i'o!;eii  store. 1  in  the  lixcr  is  iMadually  <j:iveii  up  to  tlie  hlooil  of 
the  hepatie  mmii  at  sin-h  a  rale  as  to  tiiainiain  in  the  lilood  of  the  s\  s- 
temic  eireuhilioii  a  more  or  less  eoiistant  pe  nta'je  of  'jlueose.  I  nder 
onlinary  conditions  this  process  of  <////.  .i./<  yi/i/i/v/v  is  re!ati\i-ly  slow,  hut 

when    the   i-einiireiiieiits   of   the   .n-uaiiisiii    I'or    fuel    I mi'    increased,   as 

<luriii^^  miiM-uIar  e\ereisc.  it  hecoiiies  very  rapid.  The  Mlyc.e.r.'nic  func- 
tion of  the  liver  appears  therefoi-i'  to  exist,  in  part  at  least,  for  the 
i.iirpose  of  iirevcnlinv'  the  ll Hiii;-  of  the  liloo.I  of  the  systemic  circu- 
lation with  exi-ess  of  sUL'ar  diirinu  ahscnption  from  the  intes!iiu>  ainl  ol 
mainfainiiiL.'  the  normal  perceiitaue  at  other  tinn's.  This  function  is 
aiiMloiioiis  to  that  occurrinu'-  in  plants,  in  which  the  sni;ar  in-odiicecl  in 
the  leaves,  if  not  ininiediately  rei|uired.  is  traiispoited  to  \aiions  parts 
of  the  plant  and  there  con\crter|  into  starch,  whic-li.  when  the  iilant 
l-ecinires  it,  as  during  in-w  urowtli,  iiiav  aiiain  liei-oiiie  t  raiist'oriiied  into 
ulucose. 

The    iiu'cncy    eouxciliinr    tin'    i;1m-oi;cii    into    ulucose    is    the    diastatic 

I'll-,'.) 


'I 


w 

,-['•  i 

rr 

1  -"         - 

1    ;■  ■: 

'WL    ! 


i  1 


'   1= 


:i     -I-   If! 
H    r'  if' 


!'         !■ 


,     !^ 


Jl-   : 


(wO 


MI.TMIOI.ISM 


cn/yiiio  iihii-ii;/!  iiiis( ,  wliicli  is  pri'si'iit,  imt  cmly  i.i  tlic  livt'r  fell.  Inil 
iilsii  ill  till'  Iiliiiiil  Miicl  Iviiipii.  Il  is  ii  (liniciill  iiintti'i"  to  cxplfiin  v  li.\ 
L'l.vciist'il  sliiiiilil  lie  iilili'  til  exist  iit  ;ili  in  tlic  liver  cells  in  tlie  ))i'e-ienee 
lit'  tliis  |iii\verl'iil  I'll/vine.  Tlie  fulliiw  infjf  |iiissiliilities  iiiilV  lie  ciiiisiiliM'eil: 
I  1  'i'lijil  i;I_veii'rin;ise  iliies  iiiit  feiilly  exist  in  the  livinu'  lixer  ci'Ils,  Imt 
is  ;i  piistiiiiiltrni  pi'iiiliict:  Ci'l  tliflt.  jiltlmnirli  jneseiit,  i.'l>  eii<ien;iso  is  lire 
\enteil  t'roin  iietiin:  im  tlie  •_'-|\c(n:en  in  tlie  li\  in','  liver  cell  nil  Mceoiint  "( 
tlie  hitter  lieinir  |iriit eeteil  tVeiii  its  inlllleiiee  liy  eiiiiiliin;it iull  A\itli  tin' 
snstent.ieiil.-if  siilistjiiice  :  (ir  i:!^  tlnit  MMiie  elieiiiieal  siilistiiiiee  in  the  liver 
cell  jirexents  the  "ilyei  <;eiiM  e  I'ri'iTi  .-letintr  im  the  irlycd'.'en  an  anti 
ul\  eii'_'eiiase.  Since  the  reiiiii\al  nf  any  iine  ot"  these  iiihili:tin!r  indu- 
ences  W'liilil  cause  !.'1>  eiejell  ilysis  to  lieemne  excessixc.  ami  sii  liiinir 
ahiiiit  h\  pel  ulyeeniia.  it  is  iiiijinitant.  in  searehiin;  I'nr  the  pnssilile 
ca'ises  III'  this  ciiiiililinn,  tu  examine  the  e\  iilence  that  has  lieeii  hrmiLrh) 
fiiruaril  in  sii|ii)iiit  "f  each  nf  these  views. 

Au'ainst  the  \ie\v  that  'jlv  ei'm'ii.is"  is  a  |>iistnii>rteiii  pniiliict  may  he 
I'iteil  the  \er>  I'apiil  cum  eisiun  into  'jliicuse  that  ncciivs  when  y;!ycoij:en 
is  aildcil  til  livinyr  lilimd,  as  hy  injecting  sume  inln  a  \  cin.  ( >ii  aecnnnt  nl 
the  active  '.'lyciu'eiii'lv  1  ic  action  nl'  lileml.  it  has  lieen  sii'.'iresteil  that 
flnriiti;  lite  L'lyenijin  dues  nut  leei'ine  transriirined  intn  irliiciiM-  until 
after  it  has  Icin  disehar'.'ed  intn  the  liliind  I'ruin  the  liver  cell.  When 
increased     ii'sir  must   lie  mnhili/eil.  ulv chinch  i)asses  nnehanucd.  nr  per 

haps  as  SI  me  dextrine,  into  the  lil I  and  l.vmph  nl'  the  liver  capillaries 

and  l.\  iriphaties,  the  trlyciit.'enase  iit'  which  ciuiverts  it  into  irliicnse.  tin' 
coin  etsiiiM  heiiiir  so  ia|iid  that.  li\  the  time  the  lihiod  has  traveled  from 
the  lixii-  ihronuh  the  heart  and  pnlmonary  xissels  to  the  arteries,  all 
the  lil.v  co<_'en  lias  already  heconie  transformed  into  udiiciise.  I'listiiiortcm 
udx coiieiiolysis.  accordiii'j  to  this  \  iew.  is  due  to  the  opposite  occni 
reiice  the  transference  of  u:lycoi.;enase  from  the  liliiid  into  the  livei 
cell.  Snmc  facts  supi'oi  t  ill'.'  this  \\r\\  arc  as  follows:  il'  It  has  liecii 
found  that  the  amount  of  free  jiliicnse  in  the  lilood  nf  the  vena  i-a\.i 
is  sonietinies  less  than  in  that  collected  simultaneously  from  the  earolid 
artery,  C2)  After  u'ixiiii,'  certain  siilistauees,  siieji  as  phosphorus  m 
|)eploiie.  there  is  distinct  diminution  in  the  amount  of  udyeoL'en  in  tln' 
liver,  accompanied,  houcxer.  Ii.\  no  increase  in  the  aiiionnt  of  ^Ineos'- 
in  the  lilood.  And  loi  if  the  liver  of  an  animal  that  has  lieen  rendered 
dialiclic  li.N  stimulation  of  the  splanchnic  ncr\e  or  liy  puncture  of  tin 
tloor  of  the  fourth  xentriele  is  examined  microscopicall.\-,  after  staiiiiii'-' 
liy  the  carmine  iiicthod.  masses  of  stained  Klycotren  can  he  found  present 
in  tlie  capillaries  i  siiiiisoirjs'!  that  lie  amonir  the  li\er  cells. 

Accorilinvr  to  the  second  \iew.  the  trlycnjreii  is  removed  from  tic 
inllueiiee  of  the  iiit  rahepal  ie  yK cojienase  on  aeeouiit  of  its  comhinati'in 


Tin:  Mi.TAiiDT.isM  (IF  Tin;  cAunoiivpUATus 


671 


with  ii  sustciitiiiMiliir  iiiati  ri.il.  Hy  disrupt iii'.'  tliis  (•Miiihiiiiitioii  iiiid  thus 
.•x|ic(siiis.'  till'  irlyc..i.'rii  to  thi'  iictioii  »<  v'l\''"^'''iiii'"''-  u'l.v.'i.^'ciHiiysis  will 
occtii-.  Wo  niiiy  •■fill  this  the  mci'liiiuicnl  iiypothcsis  iiihI  it  (K'sorvcs 
s.-rioiis  (■(msidcriiti.Mi.  fur  it  hjis  liccn  sliowii  lliat  very  little  iiostinortfin 
crlycojrclH.lysis  (ici-iirs  in  tin-  iiita.-l  li\<T  „(  t'rosrs  in  winter, even  thnnjih 
at  this  tiini'  thf  (>ru:an  funtains  an  cxim'ss  oI'  irlyc..!.r,.|i,-  l)nt  hccnincs 
Mjarkcd  wln'n  tlif  liver  is  Inoken  down  liy  iiieehanieal  means. 

Tlie  third  view  di>i)ends  on  the  well-Uiiown  faet   that  en/yme  activities 

I me  most  iiiarki^dly  altered  liy  sliirht  ehaiiires  in  the  ehiMiiieal  nature 

of  the  environment    in   which   they   act.      Diastatie   en/ymes  are   ])artic- 

ularly    s\iseei)1ilile    to    the    reaction    M' f   tlieir    environment,    a    very 

slidit   cleuiee  of  ai'idity    favorinu'   and    a    trac f   alkalinity    markedly 

dcpressinir  their  activities.  That  -i  tendency  to  inereasing  acidity  in 
tlic  liver  cells  may  retard  the  forination  of  'jl>co<_'i'n  is  suirircsted  hy 
the  depressiiijr  effect  ]!rodni-ecI  on  the  assimilation  limit  of  snirars  liy 
administerin!,'  aciils.  and  hy  the  ohservation  that   i>.istmortem   j.dyeo;.'en- 

(ilv.sis  111 mes  marke.l  in  iiropoition  as  the  <\\h\<:  liver  lieeomes  acid  in 

reaction.  It  midit  he  thought  then  that  srlycoirenolysis  in  the  liver  cell 
could  he  set  up  liy  the  local  pnnluction  of  a  certain  amount  of  acid. 
Such  a  lilieration  of  free  acid  I'ould  hi'  hroiiulit  al»>ut  hy  a  eurtaihuent 
in  tlte  arterial  lilood  supply  of  the  hepatic  cell,  produeiiur  a  local  accu- 
mulation eitlu-r  of  earlionie  or  of  other  less  comiiletcly  oxidi/cd  ai-ids 
ie.  R.,  lactic).  It  may  Ik'  that  a.sphyxia  causes  hyi)ers.dycemia  hy  such 
a  iiiechani-in.  Vasoconstriction  and  eon.sci(Ucnt  curtailimnt  of  arti-rial 
hlood  suiiply  occurs  in  the  liver  when  the  lii'patic  nerves  are  stimulated, 
and  it  is  imssihle  that  the  ulycoireimlysis  which  is  also  si't  uj)  hy  such 
stimulation  is  dm'  to  the  apiiearance  of  acids.  The  acceleratintf  ciTect 
of  e|)inephrine  on  silycofrenolysis  miLrht  also  he  explained  as  due  to 
limitation  of  Mood  supply  on  account   of  vasoconst rii-tion. 


ill 


An 


THE  REGULATION  OF  THE  BLOOD  SUGAR  LEVEL 

The  level  at  which  the  concentration  of  sU'.'ar  in  the  systemic  hlood 
is  maint.iinei!  reinesents  the  halanee  hetweeu  two  o|>piisins;  factors:  (li 
the  consumption  of  v'liicose  hv  the  ti.ssues,  and  i  "2 )  tlie  jirouuction  of 
■rlueose  hy  the  liver.  Since  this  is  the  most  n-adily  oxidi/ahle  of  all 
the  proximate  principles  of  fiio  i  i  pap'  (i.VJ',  mus.-ular  ,ii-tivity  c.niscs 
lar}fe  (piaiitities  of  it  to  he  consumetl.  so  that  its  concentration  in  the 
hlood  teiuls  to  fall  helow  the  jihysiolotric  level,  a  tendency  which  is 
inunetliately  nset  hy  an  imrcr.M'il  ilivi-lKirjic  nf  ^-I'li-nse  from  the  liver. 
The  (piestion  thei-efore  arises  as  to  limi-  lln  iiiiisihx  or  ntlur  lissms 
transmit    Ihiir    rt  <iiiin  im  iitx   fur    iiliicn.ir    to    th(     linr.      There    are    two 


'i'i    t 


r       St 


<i7l2 


MITMinr.IsM 


I      i      ' 


II  '  n 


iiJM 


|)ll^•.illl(■  \\;i\s  liy  uliirli  tliis  cuuld  ]•<•  dcHii':  i|'  liy  iiiriiiis  ul'  ji  iic|-vi>us 
Iclli'N.    dl'    I '_' I     liy    cllJlllUi's    ill    till'    iiiiiipiisitioli    III'    till'    liliiml.    ritlllT    witll 

i("_'.ii''l  I"  till'  |ii'iiM'iil!ii.'i'  111  siiviiir  itM'll'  uf  lii'i'iiii'^i'  111'  till-  ;i|i|ii'iiiinii'i'  in 
it  111'  ilii-iiiii|iiisitiiiii  |ii-iM!in-ts  lit'  u'lnciiM'  111-  111'  siiiiic  spi'ciiil  rxrilinir 
;i'_'riil   nr  liiiriiiiiiii'. 

Ill  iinlir  III  iisi'i'i'tiiiii  till'  ii'!,iti\  ('  iiii|iiirt;iiiri'  nl'  flii'sc  iih'IIiimIs  of 
iMilli'liitinli  lii't«iin  till'  pljiri's  III'  supply  :ili(l  iIiIIIIIIkI  III"  sjluriisi'  ill  till' 
l|ii|lli;il  ,'lllilli::i.  it  is  lli'rrssiiiy  In  ili\  est  ijjiltr  tlir  riillilil  inns  Mlliji'l'  wllji'll 
III!  i'\rissi\i'  ijisi'liai'Ji'  111'  iillli'iisi'  iiccill'S  iis  ii  I'l'siilt  111'  ii\  iTstilllllliitiiiM 
111'  till'  in'i\iii|s  ciiiitiiil.  Ill-  lii'r.'iiisi'  111'  till'  |ir('si'iii'i'  ul'  iNi'iliii;^  siilistaiii'i's 

I  liiiilnm  's  I    in  till'  111 1.      'I'lir  i.'lyi'11'Ji'llii'  I'lllii'tiiill  i-aii  In'  I'M-iti'ij  tlirnll'ill 

tilt'  nil  Us  s\s|i'iii  in  a  \iirii'ty  m'  \\:i.\s  sn  as  In  priiijiicc  liy  pi'iirlv  I'l'iiii;! 
ami  irlyi-iisiiria,  'I'liis  i-mistitiiti's  mn'  I'linii  nt'  i  ijn  r'niu  iilnl  ilinhi  l< n.  In 
laliiiratnry  animals  iin'i'liaiiii'al  ii'ritatinn  ul'  tin'  iiii'iliilla  ulilmiLMta  anil 
stiiMiilatinii  111'  till'  LTi'at  splani'linii-  ni'rvi's  ai-t  in  lliis  way.     Siniilaf  stimnla- 

tinii  may  aU riii'  ninli'i'  i-i-ftain  ciiiiilitinns  in  iiiaii.     K\i'itatiiiii  as  a  I'i'stilt 

III'  i'liaiii.'1's  in  till'  riimiiii--il  ion  ul'  tin'  liluml  ran  In'  pruiliii'i'i]  I'xpi'i'inn'n- 
tally  liy  I'i'itain  ilniirs  i  plilmlii/in  ' .  oi'  liy  tin'  ri'iiiuval  ui'  i-iTtain  uf  tin' 

illlrtli'ss    ulalnls    111'    tlic    illji'i'tiun    III'    CNtiai'ts    pji'paii'il     I'lulil    tlicill.    sllrll 

as  r'|iini'pliiiiii'. 


I!::  \i 


Nerve  Control  and  the  Nervous  Forms  of  Experimental  Diabetes. 

Tin'   siiiiplist    i\|ii'iimiiital    rirnlitiiiii    «liii'li    illiist fati's    ilii'    ii'latiunsliip 

ui'twi'i'ii    till'   iiii-viiiis  sysli'iii   ami    tliu   lil 1   siiirar   is   ilnlriciil   xlltiniliitlini 

uf  Ihi    (iri'i!    ■iiiliiiirli  nil-    III  rci    in    animals   in    wliii'li.   |i_\-    pi'i'viuiis   I'l'i'ilini.' 
\\illi  rarliuli\  ilrati's.  a   lai'L'r  amuiint   ul'  !;lyi'iii;i'ii   lias  lirnn   ili'pusiti'il   in 

till'  liM-r.     I'lV  I'Naiiiinatiuii  uf  tlir  lil 1  as  it   is  ilisi-liartrnl  intii  tlm  vena 

i-a\a  I'lulil  till'  lii'patii'  \riiis.  tin'  iiiiMras.'  in  liluml  siiirar  is  vi'i-y  I'viilciil 
ill  I'rum  tl\i'  111  ti'ii  miniilrs  al'liT  tin'  lirst  a  pplii'at  ^m  ul"  tlir  si  iiiiiilils : 
liiit  it  is  nut  until  latiT  that  a  i^nn'ral  liypnulv  i-i'iiiia  liri'uiiii's  rstali- 
lislii'il.  'I'lii-  ruiii-liisiuii  MJiii-li  \\i'  iiia>  ilraw  t'l'uiii  tln'-r  fi'siilts  is  tlial 
tlir  splani'linii'  nrr\i'  I'unfaiiis  rtl'iTi'iit  lilnTs  ruiitiuninu  tin'  fatr  at 
wllirll  u|>cui;i'n  lii'i-uiiii'S  ciilix  i-rti-il  til  iilui'usi'  ill  till'  li\i'r.  TllC  ri'llti'f 
I'iniii  wliii'h  tlii'si'  lilii'is  ui-iuinati'  is  siinati'il  suiiicw  ln'ii'  in  tin-  mnliilla 
uliliiiiL:ala.  I'ur  iln-  ii-iitaliuii  tliat  is  sit  up  liy  piiiii'tiiriii'/  this  |iiiitiun  "I' 
tin  in'iNuiis  systiiii  with  a  iirnjli'  \  iilijs  ii'iilts  similar  to  tliusi'  \\irn'li 
I'ulluw  splaln-liiiii-  si  iiniilat  iuii.  This  "  i/l  i/rni/i  iiir"  nr  dliihillf  ciiilir.  as 
it  has  In'rii  calli'il.  must  lir  pruviilnl  wilh  alVrrriit  iiiiptilsi's.  Siii'h  iiii- 
pillsi's  lia\i'  iiiilii'ij  liirii  lisi-rilii'il  ill  thr  vaiiiis  iici'vi's,  Init  their  dciii 
"iisl  rat  i-'ii  is  I'',  iiti  )i;'Mi;s  an  i-,is\  inallri-  un  ai'i'uUnl  iil'  llii'  il:.st ttrhani'i' 
ill  till'  ri'spiratuiy  muM'iin'iits  ruim-iilintlv  pMuilui'i'il  liy  thr  stiMiiilatiuii. 
Till'  I'haii'/rs  that   siirli  ilisturliaiii'i's  liiiim^  almiit   in   tlir  acfiit imi   ul'  llii' 


Till.    Ml  TM'.ill.ISM    OK    Tin;    <A1I1«>I1  Vl>ll  ATIS 


673 


1,1, ,1x1  iiiiiN  ill  tlifinsrlv.s  l.,'  |•.•sl.c.ll^il.l.•  lor  the  li.vp.T'.'lyi'rinia  (sec  iiim.' 
:;:;•_>  I.  It  .mii  :i1  Inint  !«•  siiiil  tlint  wlitii  llif  rcMpinilcny  clisliiil.iiii<-.s  iii«' 
tliiiinltil  ii-.Miiist.  iis  liy  iiitnitriK-li.'iil  iiisiiniiilion  ol'  ..xyjf.-n,  vajjal  liypcr- 
..-lyn-iniii  is  niiK'li  l.'.s  i.iJiikfd.  if  iM.I  .•iitircly  iil.s.-nt.  Kilt  this  qii.'stioii 
awaits  iiicpic  tliormisili  iMVcstii.Mti<ili. 

Tlif  increased  i.'l\  <c.L'eiiiilysis  wliieli  results  t'n.iii  stiiiiiilatiim  of  tlio 
elTereiil  lil>eis'in  tlie  s|,iaiielini<-  nervi-s  may  ilepeii.l  eitlier  on  a  direct 
eonlrol   exeieised  over  llie  ■.'!>  eoireiiie   I'lmetions  nf  tlie   liepatie  cells,  or 

on    the   diseharce    into    tlie    1.1 1    ot'   some    Imniione    wliieli    excites   the 

irlycotrcnolxtic  proce-s.  It  must  fiirtliermore  not  lip  lost  s'tjht  of  tliat 
tiie  (ilyo-^renolysis  may  he  s.'condary  to  local  aspliyxial  conditions  in 
the  liver  cells  i-esultiiii:  from  vasocoiistric-tion.  From  Iheir  anatoniie 
position,  the  adrenals  ale  to  lie  thoui:ht  of  as  Die  s.piiree  of  the  horniono. 
and  evidence  that  splan<-hnic  liyp.'r^'lyc.-inia  is  due  to  hypersecretion 
from  these  i.dan.ls  has  seemed  to  he  furnished  hy  the  fact  that  after  tliey 
are  extirpated  splanchnic  stimulation  no  loii«vr  produci's  liyperulyceiuiil, 
iM'ither.  indeed,  does  |.iiiietuie  of  the  medulla.  There  is  also  no  doulit 
that  the  nervous  system.  a<-tiii!,'  hy  way  <d'  the  splaiiclmic  nerves,  does 
excrcis.'  a  control  over  the  dischartre  of  the  internal  scrctioii  of  the 
adrenal  irlands  and  that  extracts  of  the  j.dan.1.  which  wo  must  suppose 
;u-t  in  the  same  way  as  the  internal  secretion,  cause  hyi.er-rlyceinia  when 
iii.iec'ted    intravenously    ( epinephine    hyperjrlyceiiiia    and    ■.dycosuria't. 

I'.ilt  on  theoretical  trriiiiiids  alone,  certain  difliculties  immediately  pre- 
sent themselves  in  acceptiiii:  this  as  the  mecdianism  hy  which  tlie  nervous 
system  controls  the  suuai'  outi)Ut  of  the  liver,  for  if  increased  sujrav 
formation  in  the  liver  is  dcpcn<lent  on  a  disdiaru'e  of  ei-inephrino.  the 
,|Ucstion  may  he  asked  why  this  secretion  sliould  he  caused  to  traverse 
the  entire  ciicnlation  hefore   reaidiins.'  the  liver. 

Th.M'c  are.  lu'sidcs.  certain   experimental    facts  which   do  not   conform 

with  such  a  view.     Thus.  aft<r  com|ilete  seveian )f  the  hepatic  plexus 

of  nerves,  stiinnlation  of  the  splanchnic  nerve  does  not   cause  the  usual 

il,.ur f   hyperirlyceiiiia.   wh.'ieas  electric  stimulation   of  the   peripheral 

end  of  the  cut  plexus  does  cause  it.  On  the  one  hand,  therefore,  tlu'ie 
is  eviileiHM  that  stimulation  of  the  elTcrent  iii-rvc  iiath  ahove  tlie  level  of 
the  adrenals  has  no  etVeet  on   the  su^rar  production  of  the  liver  in  the 

jihsen if  these  inlands:  ami  on   the  other,  we  see  that  when  they  are 

lircseiit.  stimulation  of  the  nerve  siipjily  of  the  liver  is  clVective.  even 
thou^di  the  point  of  stimulation  is  lieyoml  them.  There  is  hut  one  con- 
clusion that  we  may  draw  naiiidy.  that  tin-  functional  intejrrity  of  the 
cfrcifUt  nervf-lihci-s  that  control  the  Ldyeoirenolytie  i>roeos«  of  the  liver 
<lepeiids  on  1h.'  presence  of  the  adrenals.  v.My  luohahly  heeause  of  the 
iiormoiie   which   the   irlands  secrete   into   the   Mood.     This  conclusion   is 


I 


m 

wn 

i ' 

'    hi 

■  ' 

i          * 

'  j 

.:|    . 

W 

l' 

I  tl:.  1 


K 1- 


!■! 


W.I 


I       i 


•;:» 


Ml  T  Midi  ISM 


cciricilioi.ili'il  liy  tlic  I'jirt  iIimI  st iiimliit ion  i>\'  the  lirpiilic  |)lrxns.  even 
witli  a  Ntinii','  cli'c'trii'  I'liiiciil.  Millie  limi'  iil'lfi-  r(iiii|»li'tc'  rgiiii>\iil  "I 
liolli  .iilriiijils.  is  iiiit  t'lilliiw  gil  liy  llic  iisiiiil  ilc'.'rrc  n!'  I'xcili'iiii'iif  df  tin' 
L'l\  i'(>!;i'ii(il\  tic   judccss. 

Tlii'si'    i\|ii'iiiii.  Ills    ilfiMiiiisir,iti'    nil    iiii|icii(iiiit     rcliitiiinsliip    lictwcrii 

flic    IICIMiiiN    i-nlitlii|,    mill    ;it    lc;i-.|    iillc    t'lillil    nt'    ||iiriiii>||c    collll'iil,    of    tllc 

MiuMf  "iitiiiil   lit'  ihc  liver.     Tiii'x    iiiiliciiti-  tli.it   wlicii  ii  siiililcii   iiicrfiisi' 

111'  liliiiirj  siiu'ill-  is  lci|l|iii'i|.  till'  iilvi'iiL'cliii'  cclitlT  sends  out  ilil|il||ses 
«liiell    liiit    Hilly    ilireelly    exeile    the    lile.ikiliiwii    iit'    !iiye(ii»eii    in    tlie    lie 

I'll'i lis,  lillt   lllsu  siiiililt.ineiiiisly    iiil|:ienee  tile  ,ii|l  eiiiils  in  sllell  II  liiJlll- 

liei-  ;is  til  |ilii(ll|ee  iiiiiii'  e|iiiie|iiililie  in  the  liliiiiil  niiil  sii  ailltllielit  the  Jli'- 
tiiili  111'  the  iiei'\  (•  iiii|i|ilse. 

We   .ire   ;is  yet   (|liite   in    the   ihirk    .is   In   the   iiieeliiiliisni    liy   wliieli    the 
nerve    iiii|iiilses   or   the    liiiriiiniie    liriiiu's   iilidiit    inere.iseil    !.'lyiM>j;»>llolysis. 
It  must  eiiiisi>t  111'  .1  lelinivjil  (if  the  inllui'liee  tliiit   |>l'e\ellts  L'l\  ('"iri'lKilysis 
fn.iii  (leeiin  iiio   in  the  niiini.il  liver,  fur  it   liiis  Iieeii  slmwii  liy  ilireet  "h 
serv.-itinii  tli;it   there  is  nil  iiii-niise  in   the  .'iiiiuiint   nf  L'tyeiiijeiijise  lireselif 
in  e\li';icts  uf  the  liver  reiiiuvdl   fiinn  iii;iliet  ie  iiniiiiiils  nver  that  present 
ill   e\tr;iets  of  the   liver  uf  lioriil.'il   iiniliiiils.      The   piissilile   initlire   uf  this 
inlliieiiee   has  alnailv    I.ei  ii   ilisciis,ei|    I  pai,'e  tiHlli.      Thi'  chaiiLri'    iiiav  emi 
sist   either  in  t\   hiuMiiinu:  of  the  eiiiiihin.'itiun  lietweeii  the  iriyeoiren   ami 
the  l)riitnplavni  uf  the  liver  cell,  nr  in  a   rcinuval  uf  the  eheiilieal   ilidllellee 
that    iir'liiiarilv    prevents   the   Hlyeiii;eiiase    frmii   att.-iekini,'   the    irlyeii<;eii. 
In  the   liiriaer  ease  the  silyeuneii   lilierati'd   frniii   its  iiniuii   with  tile  siis 
telltaeiilar  slilist.lliees   vviiiilil   either   lieeiilne  attaekeil    Iiy    the   i;lyeiit;eliase 
Itreseiit   in  the  liver  eell   itself  or  it   wiiiild  first  (if  all  iiiiu'rntf.  as  dyeo 
i:eii.    iiitd    the    liliiMil    eapillaries    .•md    there    he    attacked    by    flie    liludd 
j.dyed(_'eliase.      l^v  ideiiee    t'ur   the    piissihility    uf  the   dcclirrelice   of  sildi    a 
priieess    h.is    .ilready    heen    yiven     i  paye    (17(11.      The    eheiiiieal     elijiii!;!'    re 
iVrred  td  inider  the  secdiid  piissiliilify  iniiiht  ennsisf   in  iiii  alfciafioii   in 
the    li.v  ilrdtrcii  idii    cdncentriifidii    uf    the    li\cr    cell,    a    cliaii'fe,    linwevcr, 
which    fur  dliviiiiis   reasuns  it   is   iiiipussilile  In   iiivcsfisrafe. 

Nervous  Diabetes  in  Man,-  The  main  interest  aft.ichiiii,'  to  the  jnves 
tii;,ilidii  (if  these  nervdiis  t'diliis  iif  eNperiiiieiital  diabetes  depends  nii  flii' 
insiv;|it   vUiicli  they  alTurd  us  iiitd  the  ii;itiire  of  the  mcehanisiii  l>y  wliieh 

i(  pioinpt    Iiili/..itiiiii  df  v'liicdse   may  he  lirdiiirlit    aliout  in   tlu rmal 

a:i;inal.  Tlicie  is  also  .snine  evidence  that  a  relaf iniiship  inav  exist  he 
tvveeii  certain  (if  the  clinical  viiiiefics  df  the  disease  in  man  niid  repejited 
excifafiiiii  df  L'lyeiiu'cndlysis  lirdiiirht  .ilidiif  by  nerve  stiniulatidii.  In 
ere.'ised  irliicdse  initMiit  fniiii  the  liver  ;is  a  resul!  uf  nerve  exeitatidU 
may  be  a  nurinal  piiK-ess.  but  there  is  reasini  td  U'licve  that  frc(piciil 
fcpelitidii  (if  this  pnieess  tends  tn  induce  a  iiermaneiit  rise  in  flie  (^liicdsc 


Till      MITMliil.lsM    UK     llll,    <   XUIlMllVliUVrr 


f.T.'i 


l,.v..l    „(   llir    I.Im.hI    mImI    tli.ivlor,.    M    trn.l.'i.rv    to   .liiilMl.s.      Tli.'iv    l.»V.' 

,,,,,,„tK    .„   .■olI.Ti..,!   M.vral    f.i.-ts   uliirl,    l.„.l   v.„„.   su|.|>'"l    to   tins 

viov.    "The    tVr t    o..r..rr..|M I'    .1,; fs    in    tlioM'    piclinpos...!    l.y 

i„t„.|itii.irr  lo  nrnroli.-  ■• li'ii.h^.  or  in   tln.sr  Nxlios,.  .lailv   In.liits  rnfml 

,„„,!.  n.Txr  Mrain.  .•umI  iIm^  .■lumavarhni  of  llir  symptoms  uln.'li  is  lil«-l.v 

,.,   |'„ll.,u    wlnn   II  .liMl"Mi.'   |.:itirn1    .MMii.. s  sonir   nnvou.  -liorU.   all 

point  in  tliis  iliirclion. 

Dial.H.s   is  roninn-n    in    I n,oli\r   <  n-ihr.is  an.l    in    tin-    .Mi-tains   of 

„„  lin,.,s      llial  is.  in  nn'n  uln.  in  tlir  |hi  roitiiMn. f  tlicir  <lail.v  illlti.^s 

„,•   fmiurntl.v    put    ihmI.t  a   s-sm.   n-Tv,-   strain.      It    is  apparmtly    in- 

,.r..asin..'  in  mrn  .•nirav-Ml  in  iipations  tlnit  .Im.an.l  mmtal  .•oin-cntra- 

tion   an.l    strain,   suri,   as   in    prof..ssional    an.l    l.nsin.ss   xvori^.      Cannot.-' 
f„„,nl  L'l.vosuria  in  four  out  of  ninr  stu.lrnts  aftiT  a  sr\.'ro  .•Nau.inaln.n. 

|„it  only  in  on.'  u(  tii.'ni  aft.'r  an  .'asi.T  ,  xa.ninati..n.«     In   tin-  nrii f 

lu.Mity four  ni.'inl..-rs  of  a  fai is  f.r.ll.all  s.|na.|.  sin-'ar  was  f..un.l  pnx. 

,.nt  in"  tu.'lv.'  iniinc.liat,.|y  alLn-  a  U.'.'niy  t.'st.'.l  •.-am.'.     Anxi.'ty  an.l 

..s,.it.'.n.nt  u.'iT  nspoiiMlil.'  t.n  its  apiM-araii.T,  f.ir  llv.'  ..I'  llu'  tw.lvp 
V  I'f.'  sul.stitutcs  uli.i  .li.l  n.'t   '.'.'t   int..  tli.'  <-'ain.'. 

.\illi..u.:li  tli.sc  n.'rv..ns  .■.,n.liti..ns.  I.y  .■x.'ifni.iit  ..f  Inl.atic  -Jyc..- 
..'.•n.ilvsis.  pr...lu.-.'  at   lirst    ii.,lliin..'  iii.iiv  tlian  an   .•s.-.'>si\  .•  .lis.'liar..'.'  of 

sipjar   int..   tin'   M I     a   .•.,n.liti..n    wlii.-ii    is   fxa.-tly   ilupli.-at.'.l    in    ..ur 

taI.orat..ry  .■Ni..Miin.nts  l.y  >tininlati. f  tli.'  in'rv.'  supply  ..f  tli.'  liv.T 

l|„.ir  r..p.'.titi..H  may  -^ra.'.ially  I.m.1  L.  tl,.'  .l..v..l,,pni.nt  ..f  a  i..rinan.'n1 
fnri.i  of  liyp.Muly.'Uiia.  T..  p.-.'v.'nt  tli.'  r.'i..-titi..n  ..f  tli.'s,.  transi.'iit 
liyp.'rt'lycfiiiias  must  !..■  ..n.'  ..f  ..ur  aims  in  tli.'  t  r.'.itm.nt  ..f  .iiHy  sta.^'cs 
..f  tlic  disease. 

Altlionuh  llu-r.'  .-an  1..'  no  -lonht  that  tin'  -Iv"!:'''''"'  fiin.-ti.ni  ..f  tlio 
liv.'r  is  sul.j.'.-t  t..  n.'rve  e..ntr..l,  it  is  pr..l.al.!.'  tliat  its  eontn.l  l.y  lior- 
Mi.in.'s    is  ..f   .'.(ual    if  n..t    ui.at.T    inip..rtane,'.      Tliis   .lual    .•..nti..l    »(    a 

,_.l;,n.lular  l lianisui  is  l.y  n..  m.'ans  uni.iii.'  f..r  tlie  ..'ly.'..!.'.'ni.'  funetion. 

f..r  w.'  liav.'  alrea.ly  s.'en  it  t..  exist  in  th.'  ease  ..f  tlie  .jaslrie  ..'laii.ls 
ami  tlie  i.a.ier.'as.  ami   it    is  ,,r..l.al.l.'  tliat    il    als..  .'xists   in   tlie  eas..   ..f 

the   lliyr.'i.l.      li    may    \\>U    W    tliat    tli.'    n.'rx tr..l    .-f    tl..'    u'ly.-..'4.'nie 

funetion  has  t..  .1..  ..nly  uith  tln.se  transit. .ry  elian-j-s  in  su'.Mr  pro.ltie- 
ti..n  that  u..nl.l  1..'  .l.'niau.l.'.l  \'>  sii.M.'n  aeliviti.'s  ..f  m.isele.  an.l  that 
tl,,.  li„.'m..n.'  e..iitr..l  has  1..  .1..  with  tin'  tti..r.'  p.'rmati.'iil  |.r...'.'ss  of  l.iiil.l- 
insr  up  an.l  l.r.'akin.-'  .l.-wt.  ..f  i-\y«'^ru  t..  m.'.'t  the  -.'n.-ral  m.'ta!...lu' 
'.('([llirenielits   ..f   tin'   tissues. 

~  -W^^.s.  l.,,.„  ,..K,M.;  ...  o,..h..M  .1..-  ..l.-.r.,.,,....  .  v.  „  .!,...»!.  th.-  rv:,„nn:>,i,.,.<  »,  r-  ,..:,.l.- 
inniNiialU    "in  r\i   i.n  kui«. 


}  V 
i 


m 


t 


II 


!£ 


.   I' 


I 


Hi' 

t  ■ 


!■  n 


(ITii 


Mn  AIIOIISM 


HORMONE  CONTROL  AND  PERMANENT  DIABETES 


N't'i'MMis  i-M-itiiliiiM  i-iiM  I'Npliiiii  iiiiIn  I I'liiisitiifv  iiii't'i'iiM's  ill  IiIikhI  siiirar, 
Mil'  limrf  |)i'tiii;iiM'iif  )i.\  |i<'ri;lvi-riiiiiis  lifiiiir  ili'pi'iidi'iit  ii|iiiii  miiiii>  ills 
Imliiiiii'i'  ill  till'  liMiiiMiiu'  i'iimiidI  III"  ciiilicilix (Initf  iitiliziitiini.     Tlii<t  <lis- 

tlll'liilllri'  is  il  MIIH'll  llinii'  M'liiiils  illYilil'  tilllll  tlllll  pl'ii<lll*'i'c|  liv  IIITX  nils 
cxcililliiiii.      Ill    tin'    IjitttT    I'iisc    tilt'    li>  pi'rirl.M'i'Iliiil    ri'iiscs    w  liclii'ViT    nil 

iif  till'   irl.M'MHrli   stcilcs  III'   llir    li\cr   IllINc   I II    ••\lillllstci| ;    W  llcriMIM   II   ilis 

turliiilM'i'    ill    llic    Ihil'liiniK'    r(iiiliiil,    Ix'siiics    railsiiiit    IIS    its    lilst    step    il 

lii'i'jiUiliivMi  of  nil  till'  ii\  iiiliilili'  ul\  i'iii.'1'M,  ijiK's  nil  til  <'iiiis('  a  |>riKliii'ti"ii 

of  sUlfJIf  "lit  111'  pliili'iii  A  piciccxs  nt'  itliirKiirou't'lirsis  (  llcw  I'lil'iiiatinli 
iif  irllli'"Si'l    IktiiIIH'S  slipi'l'llchli'ij    nil    mil'   nl'   U'lvcnirt  iml\  sis. 

Til  iisciTtiiiii  Ili4  imtiiri  of  llii-i  liiiriiiiini  iiiiil  tlii'  iiii'i'liaiiisiii  nt'  it^ 
(ictinli  liiis  Iici'li  I  111'  nlijrct  nl'  liin>t  nl'  till'  Icsi'aiflirs  nil  tlinsi'  I'nitiis  nl' 
ilialirtis  piiiiliiiTil  liy  cliaiiKi's  in  riTtaiii  nt'  tlii'  illictli'ss  uliuids.  Tin' 
I'nilnuiiiii  pns>iliiliti<'s  niav  In'  I'liiisiilni'ii:  i  I  i  that  it  is  llio  i-nncnitralinii 

nf  i:lllrns('  ill  tlu'  lilniiil  ;  I  "i  (  tlli'lt  it  is  till'  pli'srliri'  ill  till'  liliinil  nf  ili'i'niii 
pnsilinM  Jiln<]lH'ts  nt'  •.'lllcnsi';  l  :!  i  that  it  is  ilni'  tn  a  llnlliinlli'  ]>rn(lui'i'i  I 
finin  siiiiH'  iliH'tli'ss  u'laiiil,  Uciliiiililiir  thr  tii'st  nf  tlii'sc  ]inssiliilit ii's.  il 
is  sii|)pn»(il  that  the  iiii'i'lianisiii  iiiviihi'il  in  fhr  ml jiistiiH'nt  iH'twcoii  tin' 
lilnnil  siiirar  ami  tin'  iilvi'mjinii'  riiiii'tinn  is  mir  r\plii'alili'  nii  tin*  liasis 
nl"  till'  law  nf  mass  ai'tinii  ;  liailicly.  that  irl.\  rnjri'll  liffHiU's  cnlivcltril 
iiitn  u'lurnM'  «  hciii'x  I'f  the  MikkI  tinw  iiiir  tn  till'  livcr  cnntaiiis  li'ss  than 
its  linriiial  fniici'iitnitiiili  i>\  yliii-nNi'.  ainl  cniiviVM'ly,  wlu'll  this  liliiml 
cnntaiiis  an  i'M'i'ss  nl"  liliirnsi'.  as  ijiiiiiiv;  MliMiiptiiiii.  a  irlycnsirii-lillililiii'j 
pinri'ss  takt'S  ))lai'i'.  Althnllu'li  tlifir  ran  In'  littli'  ilnlllit  that  tin-  iirncoss 
of  i.'lyci)i;i'ii    I'nniiatinii   nr  ili'striii'tinii    will    i|i'|>i'iii|   tn   a   I'l'i'tain   cxfi'iit 

lipnii    till'    ailinllllt    nf    ylllinsr    piTsi'llt     ill    the    lilnnil    llnw  iliif    to    tlir    livrl 

I'i'lls,  \  I't  it  is  iiiipnssilili'  that  \h\>  r.iii  In'  an  iiii|iiirtaiit  iih'Miis  in  tin' 
cniitiiil  that  I'xists  lu'twfcii  sii'_'ai'  piuiliii-tinn  liy  the  liviM-  ami  sii'.'ar 
cipiisumptinii  |py  till'  tissiirs,  litTaii^i'  till'  sii'.'ar  that  is  aihlnl  In  the  itmlal 
Mniiil  iliiiiiiy  alisniptinii  wniilil  mask  any  ili'plctinii  caiisi'il  hy  siiLrai 
I'oiiMiiiiptii'ii  ill  till'  tissues. 

Tho  Sfcnml  pnssiliility  that  tlif  hnrilinlii'  is  snilli'  (It'cnnijinsition  l)rni| 
ili't  nf  jrliirnsi'  wiiiilij  ajipiaf  tn  liaxf  Miiiii'  siippnrt.  if  we  I'niisiili'f  thi~ 
hnrniniic  tn  hi'  all  ai'iil  proiliii't  (i-arhnii  ilioxidi'  nr  lactic')  jirndiici'il  \>\ 
siijiar  mctahnlisiii.  I'nr  it  is  kimu  ii  that  an  increase  in  the  lixdrnireii  inn 
cnlicelitratinli  nf  the  lilnnil  llnwiiiy  tn  the  liver  cells  excites  a  Lrlyenireii 
nl\sis.  As  we  have  already  seen,  hnwever.  it  is  dillielilt  In  secure  \ 
leriiiieiital  exiileiiee.  in  aiiestliet  i/ed  animals  at  hast,  that  L'lycnL'i  ii 
nl.stic  activity  is  readily  excited  in  this  way. 

The   third   jinssiliility     that    snme   s|)eeili('   hormniio   may    exist    in    the 


Tin:    MKTAIIOIISM    ••>•    Tin:   (  VHIinllVimATIS 


r,77 


,., 1  ..v..il,H.  I),.-  ul    ...«.-Mnl.vti..  p. H.     is  mv-.tiuMt...l  l,v  pro.  u..m« 

.li.tnrl i Ki.itr  vnrinns  ..f  ,1,.-  ,lu..tl...H  ulan-ls    p...t..MMn.ly  t  ... 

, m.s.  ,1,..  n.ln.nnis,   ,1...   pnn.,..> -Ms  a,.  pM"..a.y.     Tl.o   inrt  I- 

..v-is...!  I1H-..U..1,   11,-  .M-v.ms tr..l.  as  «.•  l,a>..  sr...,  1o  1..'  tin.  .-aM. 

„„„  ,1,..  „.ln-,al   uian.l      Wl,,,!,..,    H    is  l.y  ti,,.  pr...luHH.n  ot   l,onn..„..s 

.lin.Hlv    Tssavv    f..r   prnp.r   .•arl-.hv.l.at..    ,n..lalM,lis,n     or  l.y    tl.,-   n- 

....val'l-roM,  .1...  M 1  of  su-l.  s,.l.st .s  as  int.-l-r..  «itl,  tins  pr ss, 

„,„,   „,..  .luHl.ss   irlan.ls   funrtionat...   is  o, r  Ih-   -nam    pn.l.l-mH  ^v.• 

liiivc  to  coiisi.l.r.  ,    1-   ,    4 

Utilization  of  Olucoie  in  Tissue..     Although  H si..'.n..n,.al  .l.alM.1..s 

i,„l. 1    l.v   -iisturl s   in    llo.    U tion   of   tl.-   .lu-tloss   ulan.ls   ,s   nt 

,i,Ht  .l,.p.M.,|..nt  on  an  nps-t  of  tl...  ..'l.v.-ou,M,i.-  fun-tion  an.l  lat-.'  of  u'lu- 

,.,„...,i,,.n..sis.  tin.  ntili/ation  of  d -  in  tin.  lissnrs  ultin,a.-ly  I o.n.-s 

i„,,.,,C,,,,.l  „i,l,.     It  is  iln.r..fon.  important  tln.t  v .■  slnn.l.    -Iivn-ss  for  a 

„.,.nn...t    t nsi.!,.r   Inirtly    >^l,at    is   Knoun    r....'Mnlin.'    tin.    pro.M.ss   l.y 

„l,i..h   snuar  1 nns   utili/.-l   in   .1 -.anisn..     Tl.at    uhn;os..   1 np 

„s...|  np  l.v  a..tiv,.  n,us,.l..  tln.n.  ..an  l-  no  .lonl.t,     'Ihus.  ,1   .In.  n.ns..l..s 

.,,• loiiin  tin.  fro..^  an-  t..ta.n/r.l.  tl„.  ..l>.o..n. t.-nt,  n.n.i.an.-l  «.tl. 

that  of  tin-  othrr  L'tr.  «ill  I"'  f""'"!  '-'  '"'  ■li"'"""'''""!- 

\,  tirst  si.,1,1  it  n.iu'l.t  ai.i...ar  that  ih..  rasirsl  ^^ay  to  stu-lv   tl,.-  nt.l./a- 

,i ,f  ..'lu.-os.-  in   th-  nn.s.-l.-s  W....1.1  1..-  t M.par.-   its  ..o,n-ntn,t.ons 

i„  ,h..  l.l..o.i  llouin«  to  an.l .i-U'  from  tin-  n.us.-l.-.     Tin-  n.us.-l.-  Iln.t 

hns  1....-M  nn.st  s ssfuliy  .-n,l.lo>..cl  in  >t.,.li.s  ..f  this  Uin.l  has  l..M-n  tin- 

h..,-,rt      Son..-  v..ars  a-.'o  Starlin-.'  an.l  Knoulton  '  .-sa.nin.-.l  tl nsu.np- 

,■„.„    .,f   suuar   l.v    th..   ..s.-is...!    .nau.nn.lian    In-ait.   an.l    in    th.-.r   ..ari.-r 

,.M...nnn...ts  nn-.i   t.,  W  al.l.-   to  sln.u    that    tin-  .xt.-nt   t,.   whn-h   th.s 

....nsumpti....    nrn.l    was   4    .niliiununs    p..r    10.)    .,an,s    h..art    n,ns.-l.. 

,„.,.  h..ur.     A  nn.n-  thorou-.'h  r.p.-tition  of  ih.-s.-  ..Np..nn..-nts  lat.-r  l.y  1  at- 
,..rs..n  a.nl  S.arlin.      shou..,l.  how.-v..r.  that   tin-  r..sults  .-an   tnnusl,  no 

,.rit.-ri..n  ..f  tho  a.-tual  sun.p.i f  '-'Iu.om-  l.y  th-  tiss.n-  on  a.-.-.n.nt 

„,-  ,h..    fa.-t   that    th.-   tissn.-   its.-lf   n.ay   stor-    away    h-u- .ant<t..-s   of 

.•,.,rl...hv.lra1.-  in  an  innis.-.l  stat.-     i.-.  as  ..rlv.-....'..n, 

<.,h..r  inv..Mi._'ators  hav..  thon^'hl  to  stu.h  tl„.  ntili/alion  of  ..'l,l.-..s.. 
l.v  ..l.s,-rvin«  tin.  rat,-  at  uhi.h  it  .lisa|.i..-ars  fi-m  .Iravvn  l.h.o.l  k..pf  .n 
;  st.-ril,-  .-...nliti....  at  l.-.ly  t.-.np«'' ■'""•-'  f'"'  ^""""  '"""■'  "''*'''"  '''"'*"• 
This  pr...Tss  is  .-all.'.!  .,h,n.h,s;..  a.nl  it  has  1,.-."m  assnnn-.l  that  tin-  l..-oooss 
is  similar  to  that  ^^l.i.•l.  o.-.-nrs  in  tin-  tissn.-s  tln.ms.-lv.s  -  an  assn.nptn.... 
h.,w,-v.-r.   f.,r  ^^hi.•l.   th.-r,-   is  ,h.   war.anty,      ln,l.-...l.   it    may    r,-a.l.ly   b- 

sho«n  that   th.-  s:lv,.olvsis  nrrin-.'  in  Mon,!   has  v.-.-y   litlh-   if  anytlnn-^ 

,.,  .i,.  „iu,  ,h..  nliii/atiou  of  sr  -.v  \u  th-  ti-n-s.  for  it  has  1.-.  fo.nnl 
that    u'ln-.s-   .lisapiuars    f.-om   ..  ,nvn   l-h.-a   vt-ry   sh.wly   ...a-.-d   nnI..- 


?  1 


678 


Ml.TAlidl.ISM 


ii!l 


i 

1  i^^^ 

|:' 

1 

<    • 

i 


<'i.iii|i;n<'il  Willi  tlif  i;it<'  ;i1  wliicli  it  disiipix'iirs  iVnni  tlic  Mood  of  jiiiiniiils 
in  wliicli  till'  .•Klilitioii  of  ".'lii'-osc  from  tlic  liver  li;is  lifcii  iircvi'iitcd  Kv 
iriuoxill   of  tllis  \  iscils   (  .M;iclro(|  I,-' 

A  lliird  iiuIIhuI  for  stiidyiii<r  tin-  utiliziitioii  of  u'lncosc  i-onsisls  in 
olisciviiiir  till'  risi>ir<il,irii  i  .n-lnniiii  of  .-ininiiils.  In  iionii;il  jininuils  tlir 
in.ji'i'tioii  (if  L'liirosc  ciiusi's  iiii  ini-i'Oiis(>  in  tin'  I'.irlion-ilioxiili'  cxi-ri'tioii 
;iiid  .1  lis,'  ill  till'  ri'siiir.-itoiy  i|iiotii'iit.  wliii-li  it  will  In"  n'mi'inliiTi'd  is 
il  liitio  ('Viiii'ssinir  tlic  ii'l.-itioiisliii)  Intwi'i'ii  tin-  iunount  of  i-,iiIioii  dioxidi' 
ONiMi'tn]  ;ind  of  till'  iiNviicn  n't:iini'd  in  tin'  or!;;inisiii.  Wlii'ii  i'iirl)oli\ - 
dr.-itc  is  iiiidiT'.'-oiiiu'-  i-oiiiliiistio!i.  tlir  iiiiotifiit  is  iirMiIy  1,  w  Iiiti'.-is  witli 
lli:it  of  proli'in  it  is  .ilmiit  (t.T  (  sn'  ii.iiri'  ."47  i.  I'.y  oliscrx  iin.'  tlii'  <|i|otii'nt 
nnilt'i-  irivi'ii  I'oiiilitions  oiii>  i-jin  I'oiiipnti'  tin'  ))ro|iintions  of  I'lirlioliydiatc 
;iiid  of  f;it  ;in,l  pi-oti'in  tlijit  Jiri'  nndiT'.'oiin.'  itn't:iliolisiii.  In  tlif  Ininds 
of  Miii'liii  nnd  otinis,-'  tliis  iiirtliod  luis  jnovi'd  of  soiiif  v.-ilnc  in  srttlinu 
I'l'i't.iin  (|iii'stiiiiis  coni-i'min!.'  tin'  ntiliz.-ilioii  of  irlnrosi'  in  nornnil  find 
dijilii'tii'  jiiiiiiiiils;  Imt  till'  ii'siilts  imist  hi'  intiTpritcd  witli  >:i-i>;if  cjui'  on 
iiivonnt  of  ili(>  f.ii't  tli.it  ti'iiii>i)i;iiy  idiiui'^i's  ill  tin'  Idood  iiniy  i-.'insi'  ;i 
•rrcatiT  or  a  liss  I'Niuilsion  of  cailion  dioxidi'  from  it.  Tims,  if  ai'iiK 
a|>pi'ar  in  tin-  Moo,!,  liu.y  will  dislodi;.'  rarlion  dioxide  from  if,  and 
apparently  cause  the  respiratory  i|nolieiit  to  rise.  Alkalies,  on  the  other 
hand,  apparently  cause  it  temporarily  to  fall,  and  niiless  the  ohserva 
tioiis  ai'c  done  o\er  a  loin,'  period  of  time  and  with  -.'reat  care,  faiiltv 
conclusions  ai-c  xery  apt  to  he  drawn   from  the  results. 

Diabetes  and  the  Ductless  Glands 

We  are  now    m  a  position   to  consider  the  forms  of  experimental   dia- 
hetes   pr.Mliiced   li,\    i list  11  il i.'iiices   ill   the  ductless   ulands. 

Relationship  of  the  Pancreas  to  Sugar  Metabolism.  In  no  othci'  of 
the  ni.iny  caiis,.s  of  dialietes  has  L'reater  interest  heen  slmwn  than  in 
that  due  to  distiirlpaiiee  ill  the  iiaiiereatic  fiiiietioii.  Miiiiy  of  the  e.irlier 
clinicians  that  followed  i;is,.s  nf  di.ilietes  inellitiis  into  the  |>ostiiiortem 
roiiiii.  noted  that  delinite  inorhid  ehaiiu'cs  in  the  |ianereas  were  a  fre 
ipicnt  aecuiup.inimciit  of  the  dise.ise.  I'lompted  hy  these  ohservations. 
several  in\e,tii;:itors  attempted  experinicntal  cxtir])atioii  o!'  the  udaiid. 
leit  did  not  succeed  in  prodncin-j  udyeosnria  in  the  few  animals  that 
siir\i\id  the  operation.  'I'licir  failure,  no  doliht,  reslllled  from  iiieom 
Iilete  exiiipalioii.     To  re, luce  the  severity  of  the  operjitioii.  Clande  I'er- 

Hard  ilijcetcil  oil  into  the  iiaiiereatic  .lt|,-t.  and  ticl  it;  Init   he  s -c'd,',! 

in  kecpiii;;  ,,i,|y  two  ilo-is  ;ili\e  I'or  any  Icnjilh  of  time,  and  these  di,| 
not  cxhiliit  e-lM'osiiria.  Neither  were  other  invcsti'.'atois  that  .-idopled 
siitiil.'ir  !ii.'!  I:, ..■!•-  ;;ny  u\"-'  -neee-^f.il.  It  hniked  ;is  if  the  pancreas  h;id 
very  little  to  do  with  tl ;iuse  of  diahetes.     In  the  vear  |ss!t  Mjnkouski 


Tin;  viTAiioi.isM  OF  Tin;  cAnnonYHUATrs 


(.70 


,,,.1  V...,  M.-riMu'  in  (i.-.-n.ny.  .Mid  .1..  Don.ii.i.-is  in    Italy,  l.y  tlmnmirl. 
■..xtin,Mti...i  of  tl-  u'lnn.i.  su.-.M.,.,l...l  i„  i.nMlu.Mnu'  in  .I--'s  m  unnk.-l  an. 

,„.,sist.-nt    i;lvn.snria.  : ..ui'ani.Ml  l.y  many   ..f  Hm'  ..tl.-r  sy,ni.t...ns  ot 

,li:,l„.tcs.     Ttie  first   Um>  antli-.is  atltil.ut..[  ll..'  .•..n.rni..n  U,  .■.•n.-.val  <.t 
JIM  iiitcrnal  sc.-i'.'lii.n. 

Tl, Ufs,.  ..r  til.'  .iiah.^t.s  tliiis  ,.r...lu.'..l  i>,  h..«.'N.T,  ^ "l.al  .lilT.|r- 

,,,„  ,•,.„,.  tliat  Msuallv  ol.MTV...!  in  ..lan.  It  is  ,.Mn -inly  a.-ut.'  f.-n.  tlio 
start  tl...  <i:  N  rati,,  l-.^in.^'  I  ::!.(;  (s....  i-a-.'  MV .  an.l  it  is  MMa.M...i,.i.an,...l 
l,v  anv  ..r  tlu-  .-lassi,.,!  sy,Mi.t..n,s  so....  in  tl,.-  -•lini.'al  .-.n.litu.n,  Kxi'.^n- 
„;,,„,,;,  p,„M-n.ati.-  .lial...1..s  .-an.  Im-wcv.'.  W  nia-l.'  L-  sininlal..  v.M-y  -•l.-s.'ly 
11,..  ,lis,..s..  in  man.  Tl.is  was  first  of  all  ,l,..M..ns1,-at...l  l.y  San.l.m.'.vv. 
„l,o  f,.nn.l  tl.at  if  tin'  t.'r..af..r  part  of  tli-'  l.an.-ivas  was  ninov.-.l.  tlH' 
,„i„„ls   for  s..,„..   .....nths.   if  at    all.   w.mv   only   ......asi..nally   .^ly.M.sunc. 

1„„  l„t,.r  iM-a.n.-  n,...-..  an.l  nn.r,.  fr..,|,i.M.tly  so.  until  at  last  tl. n.l.t.on 

tv,.i,-ai  ..f  .•o.Mi.l.'t..  ,.an.M-.-at.M-t.....y  stn.-Mv...,..!.  Sii.nlar  n'sults  l.av.' 
,;,„,,.  ,,,M.ntlv  l...cn  ol.tain...l  l.y  Thin.L.ix  an.l  Ja.-.l.,  in  Kran.-...  an.l  hy 
\)lrn  in  tl.is  .-..untrv.  Tl,.-.-  inv..sti.^'at,.,s  point  ....1  tl.at  .l.n.'r..|,t  v.>- 
si.lts  arc  t..  1.."  .■xi..--t.M  a-M.-nlin-.-  to  ul,.'tl,.-r  tl,-  p..,tion  ..f  pan.Mvas 
wl,i..l,  is  ''ft  .1...S.  ..r  .l...'s  n..i,  ivn.ain  in  .•onn.M-ti..n  with  tl,.-  .l,....l.-nal 
,l,„.t      \-       n  tliis  .ln.-1   is  li.-'at.-.l.  atn-i.l.y   of  any  r.-n,na„t  of  ,,an<-,-.-as 

,I,,„  is  l..ft  is  l,o,M,.l  t... nr.  an.l  tl,is  is  nsso.-iat.-.l  witl>  rapi.l  .-ma.-ia- 

,;.,„  .,,-  „,,  ,„i,„,l.  ,li.  ..fs  an.l  -l.-atl,,  \Vl,.-n  tl,.-  n-.nnant  s>.rr..,n,(ls  a 
still  ,.at.-nt  -In.t.  a  .-..n.liti.-n  mn.-l,  nmn-  .l.s.-ly  sin.nlati...,'  .lial-.-t.-s  .n 
„„„  is  lik.-lv  t..  l...,.,.,,,.-  ,l.-v,-lop.-.l  O.H-.  .,a.„.-ly.  in  wl,i.-li  tl,.-ro  ,s,  ..r 
so,„.<  in..i,tl,s  f.,ll..^^i.,tr  tl,--  ..|..-rat ion.  a  n,.uv  .,r  l.-ss  ni.M  .liaI..L-s, 
wl.icli.  li..w.'v.-i-,  nsiially  pass.-s  lat.-r  int..  tl,.-  fatal  ty]..-. 

It   is    of  .•-..M-s.-.  .lifli.-.ilt    t..  Stat.-   a<-,-,irat..ly   wl,at    pr..i...rti..i,   of  tl,.- 
nan.-r.-as  ,n,>st  1,.-  l.-ft  in  ..r.l.-r  tliaf  th-  abov.-  .l.-vriL.-.l  .-..n.liti.-n  may  supor- 
yruo      I..-aviM.'   a    r.-mnan1    am<.u..tiM-   t..    fr..n,    on.--fiftl,    1-    on.-.-ii-'l.tli 
„f  tl,.>  .-„tir,<  ..'lan.l  is  .•.,mm..oly   f..llow.-.l   l.y  a   mil.l  .lial,.-t.-s.   wl,.T.-as 
if  n„lv  ..n.-.nintl.  ..r  l.-ss  is  l.-ft.  a   rapi.Uy   fatal   typ.-  -l.-v.-l.-ps.     .\s  n, 
,|i„i,.al  ,.xp.-ri.-n.-.-.  tl,.-  .listin-.n.isl.i,,..-  f.-atnn-  L.-tw.-,-,,  tl,.-  m.l.l  an.l  tl,.- 
s..v.-,o  tvp.-s  ..f  .-xp.-rim.-ntal  pan.-n-ati,-  .lial..-t,-s  is  tl,.-  t..l,-,-a,,...-  f.-wanl 
.,,rl...l,v;irat,.s.     In  tl,.-  mil.l   f-m,.  n-  i:ly.-..s,„ia  .l-v.-lops  nnl.-ss  .-a'-l-o- 
l.v.lrato  f.....l  is  tak.-n;  in  tl,.-  s.-v.-r.-  f..rm.  it  is  p,vs,-nt  w1,.m,  ll,.>  .In-f^  -s 
..;„„pos.-.l   .-nti.-.-lv   ..f  tl.-sl,.     It    is   tl.us  slo.w.,   tl.at    "l.y    r.-nn-val    ot    a 
snital.l.-   p,..p..rti..n   ..f  tl,.-  pa.,.-,-.-as.   it    is   ,...ssil,l.-   t..   l..in«   an   animal 
t.,  ,1,.  v.-run-  ..f  .r,al..t.-s.  y.-t  1..  know  that  th-  aniu,al  vvill  n.-v.-r  ..f  ,ts.-  I 
,„,..„„„.  ,,i,,,,,i.,     .     .      .     Sn.-h   animals.   th.-,vfo.v.   .-..nstituto   valnal.  .- 
t,-st  ohi.-.-ls  f..r  .iu.k'in--'  the  .-nV.-ts  ..f  xari..ns  a..M-M.-u-s  with   r.-sp.-.-t   1o 
,li-,h.-t«-s"     (  Ml-n'-^      It  lh.-r.-f..r.-  h.-.-.-m.-s  th.'..r.-t i.-ally  p..ssil.l.'  t..  m- 
v.'sti.'at.'   on  tl,.-  ......  han.l,  oth.-r  .■on.liti..ns  wiii.-h  will  havo  an  intln.-n.-e 


m 


li    fit 


I' 


M 


ifi 


1.        5 


■■    i; 


»'s 


lt;    ■      i 


mi 


680 


mi:taii()I,ism 


similar  t<i   iciimval   of  inoir  of  tlic  •.'l"iii<l.  or,   i>ii   tlic  dtlicr.  cinKlitiiins 

wliii'h  iniu'lil   pifvcnt   tlic  iiiriilfiK t'  iliiilictcs,  even  tluuiirli  tliis  extra 

pciitimi  iif  paiiiTcas  is  rcnioxcd. 

From  111.'  Avork  wliii-li  li,'  lias  airca.ly  cl.in.'.  All.'ii  liclicv.-s  tliat  lie  lias 

siifliciciit    cvidciicc   to   slmu    tliat    tlic   coiitiiiiicd    f liiu:   with   cxi't'ss   nf 

(•arli(ili,\(lrat('  fciml  will  surely  cuincrt  a  iniM  into  a  severe  ease,  and  in 

'" \l>eriiMeiit   lie  siieeeeiled   in   liriiijiiii!:  almnt    tlie  same   transition   tiy 

perl'orniin'i'  pniietnre  of  the  medulla     that   is,  liy  ereatinj:  an   irritative 

nervous  lesion,     ]\\   not t' the  other  means  usually  emploxcd  to  i)rodiice 

experimental  -.'lyeosuria  could  the  liorderiny;  ease  lie  made  dialietie, 
althoiiuh  one  such  animal  Iieeame  acutely  dialietie  after  liiratiire  of  the 
Iiortal  vein.  To  the  elinieal  worker  the  value  of  these  results  lies  in  the 
fact  that  they  fiiinish  eNperimental  pioof  that  a  so-calleil  lati'nt  ease 
of  dialietes  that  is,  one  that  has  a  low  tolerance  value  for  earliohy- 
drates  may  he  prevented  from  <lcvelo|iinu'  into  a  severe  case  li\  i)ropei 
control  of  the  diet.  Attempts  to  show  wliether  or  not  there  are  nnv 
eonditions  which  iiiiiiht  l)rin<r  aliout  improvements  in  animals  that  were 
.iust  ilialietii-  have  not  as  yet  heeii  sutliciently  made  to  warrant  any  con- 
<dusions  that  eoiild  help  ns  in  the  treatment  of  hiniian  cases.  The  en- 
coura'.'i'iiient  of  the  internal  pancreatic  seei-etioii  liy  diminution  of  that 
disidiarjic   into  the   intestine  may  he  of  \alue. 

The  certainty  with  which  diahetes  results  from  pancrcateetoiiiy  in  do^'s. 
as  well  as  the  fre<|uent  occtirrcn f  demonsf rahlc  lesions  in  the  pan- 
creas in  dialietes  in  man.  h'aves  no  doulit  that  this  --land  must  lie  in  somi' 
way  essential  in  the  physiologic  hreahdowii  of  earliohydrales  in  the 
normal  animal,  hut   how,  we  can   not   at   present    tell.     All   we  know  is 

that  tlie  first  (dian^e  ti cur  after  the  u:laiiil  is  removed  is  ;i  swcepiiiL' 

oiif  of  all  hut  a  tr. if  the  (.dycoL'eii  id'  the  liver,  althoii^di  the  muscles  may 

retain  theirs;  indeed,  in  the  cardiac  mu.scle  there  may  lie  more  than 
the  usual  ;imount.'~  Nor  can  any  u'lyco<ren  lie  stored  in  the  liver  when 
excess  of  earliohydrates  is  led.  Al'ter  the  ulycoo;,.!,  luis  disappeared, 
liliiconeoLrencsis  sets  in,  so  that  the  tissues  COMIC  to  melt  awa,\  into  snu'ar, 
and  all  the  symptoms  of  acute  starvation,  associated  with  certain  others 
that  aie  possilily  due  to  a  toxii-  action  of  the  excels  of  snu'ar  or  other 
aliiiormal   products  in   the  lilood,   make  their  appearance. 

So  fai  it  llli'_'Ilt  lie  pi'rillissiliie  to  eoiisidcr  an  ciM't-prodnetioll  of  irju- 
cose  as  the  cause  of  the  !iyper<.'lycemia  of  paiicreati.'  iliahetes.  just  as 
we  have  v|.,.n  it  to  lie  of  these  foi'iiis  of  hypciud  V  cemia  that  are  due  to 
slininlalion  of  the  nervous  system;  Iml  this  can  not  lie  the  cise,  for 
another  \r\y  definite  alinormalit\  in  nietaliolism  liecomcs  evident-  - 
ii;nii'-!.\ .  ah  in.iliiiit  \  of  Ihr  tissues  to  liurn  siiiiar.  This  fact  is  asi-cr- 
tained    liy  olisci>\invr   the   respiratory   ((Uotient.      When   s;lucose   is  added 


Tlir.    MITAlliil.lSM    OF    Tin:    CAItlloIlYPUATKS 


681 


,„  tI,o  Mo.mI  in  11,.-  caM-  of  n  .MMnplrfly  .lii.lM.ti.-  MiiiMinl.  ....  rliMUirc  or- 
(Miis  ill  tlio  (|noti('iif. 

T1..MV    an',    thrn-rnic.    tu..    rssnitinl    ,listiiilian<-.-s     of     carl.ol.y.lratc 

taliolis.n  in  pan.-rratic  ,|i;il„.1rs    .ov.T,,nMl..HioM   of  si.-ar  an.l  al.oli- 

,io„  of  tlo-  al.ilitv  of  111-  tissu.'s  to  us.,  it.  It  l..'.-oni.'s  in.i.ortant  f..r  us 
t„  s...-  wh.'tli.'r  tl...  tissn.'s  ..xhihit  tliis  inalnlity  to  use  su-^'ar  wli.-n  they 
•ur  is..lat.'<l  I'n.iii  tlic  animal:  f..r  if  tlo^y  sli.ml.l,  a  inu.'l.  i.k.it  s,-nivlnn« 
invcsti.Mtion  ..f  111.'  .'ss,.ntii,i  .-aus,'  „f  tti-ir  inability  \voul.l  l.o  poss.l.l.^ 
,|,,,„  is  tin-  .-as,,  wli.'n  tli.'v  ai'.-  fun.-ti..nin-  al-n..'  will.  tl..>  oil..'.'  organs 
;„i<l  tissii.'s.    Th.'  .'arli.'r  .'X|MTin..'nts  of  i/pin.'  an.l  Lis  i.n|.ils.  wl.i.-l.  s.M-m.'.l 

to   sln.w    tl.at    .lialM.ti.'    M 1    -li-l    i.ol    Im-ss.'ss    11..-    -ly.-olyti.-    imwrr   ..I 

„„,,„,,1  l,l,„Mh  an.l  tl...s..  of  (•..l.nli.i.n.  f.-o.n  ^^Uu■h  it  was  -l...!.'.!  Il.at 

,„iNt...-.'s  of  tl..-..xi.n'ss.',l  jni.-.'s  .>f  ni..s.-l,.  Miv...'.  an.l  pan.-.'.-as.  altl...nj.'!, 
„,,lina.ily  .L'slnnin-.'  uh..'.-.  fail''!  to  .1..  s..  ^^l..•n  tl...y  ^^>^u^  tak.'U  f.-.un 
;,  ,liiil..'li<-  animal,  ai.'  n.'w  kn..\v.i  to  1..-  .v.'on.'ons. 

Tl,.'  failnr.'  t,.  sliow  m  .l.|..-.'ssi..n  ..f  dy.-olyti.-  v»^^r^■  l.y  tlu-s..  m.'tl.o.ls 
p,,„„l.t.'.l  K.,..»lt.m  an.l  Staili..'-'''  L-  inv.'sti-at.-  tl.c  .|...'sti..n   wl,.'tl..'.' 
nny  .lilT.'.'.'i..-.'  is  .'vi.l.'.it  i..  tl..'  .at,,  will.  «l.i<-l.  irln.-oM-  .lisMpp.-ais  tV.m. 
;,  ,'niNt.M-.'  ..f  M.M..1  an.l  s^li.,.-  s..l..tion  .i-.'.l  t..  i...i-r.is..  a  li.'a.t  ....tsi.l.. 

,1„.  I..„lv.  M.-.-o.-.lini:  t..  ^^!..■tl..•.•  ll..'  li-arl   "as  f.-o.n  a   .....'..lal  ...'  a   -l.a- 

|,Hi,.  .1...'      In  til.'  liist  s.n.s  of  ..l.s.',vati..ns  «l,i.-l.  tli.'s.-  w...k.'.-s  i.ia.l.". 
it  was  tlm..d't  tliat  tl...  .i.n.i.al  b.'a.t  ..s.'.l  -h.-os.'  at  tli-  .'at.-  of  ahout 

4  n.-'   p.T  1<>t»  i-'i...  <'!'  l.'-a.l  snl.stan.'.'  p.'.'  ...i..ut.':  "L.'.'.'as  tl.at  ..t  a  .l.a- 
iM'tii^   (.l..pa...Mvati/.,'.l.    at.imal    .is.'.l   I--  H.an    1    ...'-'.      If  s'"'''   ^"••'<'"-' 
,li(T.'r.>...'..s   i..   11...    fat..   ..f  M.ira.'   ......s.m.i.tio.i    --....Ll    ...ak.'   tl..'...s..1v.'s 

,„,.„.ilVst  for  so  .-.'lativ.'ly  small  a  ...ass  .,f  n.,is,M.lar  t'ss...-  as  tliat  ..f  tl..' 
l,..;nt.  it  is  p.'nnissil.l.'  t..  ass,.....'  tl.at   a  .......li   n....r  st.ikin-,'  -l.tT.'.'...,.'.' 

,„„1,1  1...  ,l,.,n....st.at.'.!  «l..'i.  tl..'  p.'.f..si....  Ih.i.l  is  .iia.l.'  t..  t.av.'.'s.-  all 

,„■  pra.'ti.'allv  all  of  tli.'  sk.'l.'tal  .in.s.-l.'s,  :.s  xv.'ll  as  tin-  li.-a.t,     Vov  tli.s 

,„„.,,.,s,.  an  ..vis.'...at,'.I  a.ii.nal  ...ay  1 mpL.y.'.l     that   is.  ......  in  ^vkn-l. 

,1„.  Ml,.l..mi..al   vis.-.'.'a  a.-,'  .■..ni.iv.'.l  aft.'.-  li-ati.  ..  ...   tl lia.-  axis  an.l 

n„.s.'nt.'.'i.-  ai-t.Ti.'s.  a...l  tl,..  liv,',-  is  ..linii..at...l  l.y  ...ass  r,.,:ati.m  ..f  .ts 
l.,l,os.     rsi..u'  s.l.-l,  |'.-.'pa.ali....s.  I{.  (I.   I'.ar.-.'  an.l  .Ma-'Lo-k  '  f-wi.-l  tl.at 

tl,,.   .'at,'   at    wki.-l.   ..'1...-..S..   .r.sai,i..'a.s   tV.m,    ll..'   1-1 1.   altl.....;:!.    v.-fy 

iii',.!:iilai'.  is  i..  no  way  .linV.-.'..t  in  .-onipl.t.'ly  .lialM'ti.-  as  ...nni.a.'.'.l 
uitli  .....mal  .L.L's.  Tl..'y  «.'.■<■  tlms  n..al.l.'  to  ,•,,.. liim  any  ..f  K....wlto., 
,•11, .1  Sti.flinsr's  .'arli.'.  ,-o.„-l..sions.  I'i.tt, ■.-..,  ,in.l  Stai'li..'.'  snl,s,'.p..^..tly 
p„int.'.l  out  tl.at  a  s.'.io.is  .■,•,■...•  was  involv.'.l  i..  tl..'  .'a.li.'i'  p...f..si.... 
,'xp,'.i....'nts.   pai'tlv   ,..,    a,-,'......t     <■(    a     ,,'..ia.k.-.l.','    I..it     i.T."-'.ila.'   .!.sa|.- 

p,..n.n<' •   ::h..-os,'    f.-u'   H'"   !.i...:s.   :u.,l    ..a.Hy    1 a..s..   tl..-   -lialx't..- 

!,..art    n.av   .'....tain   a   .'....si.l.'.al.l.'  .'X.'.ss  .,f   v-ly.-.,};.'...    ffom   xvl....l.   .ts 


!  .;■  .  1  t 


^i      r 


>;,n  h 


!i  ' ';: 


I    i 


:,■.'   ti 


KS:!       '   1 


|i  ■■ 


II J  i 


r.s-j 


mi;t.mi()I,ism 


(liMiiiiiids  t'i>f  .siii.'.ii-  ni.iy  111'  met  willioul  ciilliiiy:  on  tluil  oi'  tlic  ])i'i-riisiiiii 
llniil. 

Ill  ^|iitc  iif  till'  I'.iiluiT  til  sliiiw  Unit  till'  iMiliiti'd  tissiii's  nf  (li;il)i'tic 
.iniiiiiils  liiiM'  ii  jowiT  yliii',is('-riiii>liiiiiiii:  liiiuiT  tliilli  tlmsi'  (if  llnniiii! 
,i!iiiii;ils.  it  JN  iiii|iiiitiiiit  frnin  :\  ]ii;iilii-;il  Nl^iinlpiiiiit  tli;it  \vc  slionlil 
kiiiiw  siii'irtliiiiL'  fi".Mfililiy  //m  iiiisxihli  iiiiliin  nf  Nil  ilislKrhiiiirr  wliirli 
;i  ri'iiinvjij  111'  till"  |i,iiiciiiis  ciitiiils.  Kvcii  it'  we  cinild  not  tell  cMictly 
liow  tliis  ilistnili.'iiicc  opciiiti's.  it  woiilil  In-  of  viiliic  to  know  Avlictlii'i 
it  ili'pi'iiii>  (111  tlip  ii  ?iii)\;il  from  till'  oi-Lr;inisiii  of  sonic  liorinoiic  lliiit  is 
cssi'iiti.-il   to  (•;iilioli\(liatc  ntiliz.-itioii.   for,   if  lliis  wen'  iirovcil   to  lie  tli" 

<'iis(',  el :r;i','i'iii(iit    would   I ffcri'd   to  seek    for   tlic  I'licniiciil    nature 

(if  tliis  liornioiic  so  tliat  we  iniirlit  iidniinisli-r  it  witli  the  oli.jcct  of  re 
iiioxiiii;  till'  dialii'tii-  state.  Tin'  Imi-i'  of  a  fruitful  ontcoinc  of  sndi  an 
iii\  i'sti<rati(in  is  cncoiiraLii'd  li\  tin'  siici-csv  of  rcsfarclii-s  on  diseases  ol 
(itlier  dnctli'ss  lilainls,  |)artienlail\    tlie  tli\  roid. 

Tlie    icnioxal    of   snnie    lioriiione    neeossarv    for    iniipcr    snirar    inetali 
olisni   is.  liowever.   •  \    no  nie.'i'is  tlie  onl\    way  Iiy  mIiIi-Ii   tlic  results  can 
lie  csplaiitcd.    for  we  can   assunie   tliat    llic   jiancrcas  owes   its  inflm'nec 
o\ir  siiirar   nicialiolisiii    to  snine   cliaiii.'c  oeenrriii'.'   in    the   coniposition    of 
tlic  Mood  as  tliis  circnlatcs  tluonuli   tlic   i:laiid     a   clwiniro  wliidi   is  dc 
licndeiil    on   the   inti".'iity   of  the   eland    and    not    on   any   one  enzyme   or 
hnnnoiie   whiih    it    produces.      It    is  olivions  that    the   results   of  i'cmo\al 
of  the    irlaiid    could    he   explained    ill    teiiiis   of  either   view,    and    indeed 
there  is  hut   one  e\periniciit   which  would  )>criiiit   us  to  decide  wliich  of 
them    is  eoiii'ct.      This   consists   in   secinir   wlietlier   tlie  syiiii>toms   wliieh 
lollow    pani-rcatcctoiiiy   are    reinovcd,     and     a     iinrmal     enndition     reestali 
lished.    when    means   ar.'   taken    to   supplv     the    supposed    missinir   internal 
si'i-retiiiii  to  the  oreanisiii;   if  they   should   lie,  conclusive  e\idi'nce   woiihl 
lie   furnished   that    it    is  Ky  "internal   secretion"  and   not    li.v'  "local    in 
llncni-e"  that  the  elainl   functionates. 

The  experiments  ha\e  lieeii  of  two  t.vpes:  in  the  one.  variously  ]ire 
lulled  cNlracts  of  the  inlands  Inuc  lieen  emplow'd,  and  in  the  other. 
Iiliioil  which  is  pr"sumalily  rich  in  the  inleinal  seei'etioii.  The  most 
recent  work  with  pjiin-rcatie  extracts  lias  shown  that  injection  of  i>an 
ercatic  estiacis  into  a  depancreati/ed  animal  ]Mddnces  no  clianij:e  in  tie' 
respiratory  ipiotieiit.  .■iltlmn'-di  in.iections  of  extracts  of  )'alicreas  ami 
duiiilcnnm  may  priiducc  a  temporarv  f;ill  in  the  dextrose  excretion  in 
the  urine  mi  accoiint  of  the  alkalinity  of  the  extract.  Neither  have 
cx]ierimeiits  with  lilood  transfusions  yielded  results  that  are  any  more 
s;,tisf,ietory.  In  u.:deil,ikin'^'  these  experimciit-s  it  is  of  conrsc  ;i.ssunieil 
that  the  inlern.il  secielion  is  present  in  the  lilood,  jiinj  that  if  this  lilond 
is  supplied  to  an  animal  sut1"criii<;  from  diabetes  lieeau.sc  of  the  loss  of 


1         I 


Tin:  MiTAii'Pi.isM  or  thk  cAiiisoiivimATiis 


f)83 


il.  ,.,,n.MTns.  it  ^^\\\  n-st,.r.-  it  t<.  a  ..on.li:,n..ti.-  slat.-.  Tl,.'  i:.Mu-n.l  cm- 
..lusioi.  tlmt  m.-iv  Im.  .liMVMi  tVom  lli.'  nu.M.Tous  .•(■s..inTl..'s  of  tins  u:\U\vc. 

is  tl.at   tlHMr   is  no  .'vi.l. H.^t   tin.  l,lon,l   ..I'  a   normal   animal,   ov.-n 

,vl,..n  it  is  from  111.-  pan-Mvatic  v.in.  contains  an  internal  s.-ciTtion  tl.at 
,.,„  n-storo  to  a  .liaLHi.-  animal  any  of  its  loM  imxv.r  to  ut.l./.-  .•a.l.o- 
liv.lratos.  Wlicn  tli.'  .M.'nt  of  -^Ivrosniia  alone  is  ms..1  as  tlu^  cntorion 
„t'  tin-  Stat.'  of  .■arlioliv.liate  m.'tal.olism.  s.Tions  .•irors  in  .iml-rment  arc 

,i.,),l,.  ,„  l„.  .Irawn.     Tlu'  ron-lition   of  tin.  1.1 1  sn-^-ar  an.l  tin-  .'xtcnt 

,n.l  .•liaract.T  of  11,.-  rcsi.iiatory  .•x.-lnno^.'  at.'  tl...  nn.st  rclial.lo  in.l.>x<'s. 


DIABETIC  ACIDOSIS  OR  KETOSIS 

Nature  and  Cause.     Mu.'l.  .-..nfuMon  l.as  ..xist.'.l  in  m.'.li.-al  lit.Tatun- 
nv.T  tlu'  .•o.T.'.'l  .l.'tiniti..n  ..f  a.'i.L.sis.  mainly  l..'.auso  tin'  t.-rn.  \vas  lust 

nsc.l   for  tl.o  partirnlar  vari.ty   »{  \\u n.liti..n  ..l.s.rv,.,!   n.  tlio  lat.-r 

sta.'.'s  of  .lialM't.s  m.llitns.     Tlic  a.-i-is  xvlii.'li  a.-.-nmnlat.'   n.   tlio  tissue 

llui.ls   in   tins  .lis-as..   are   a.-.M,.: ti.-   an.l   /i  ..xyl.utyrie.   an,.!   tlioy    an- 

„si,la1i..n  ,.r.Hln.-ts  of  a<-.'t..n,-.  Nvlii.-l.  is  a-.-ain  .l-riv..!   fr-.m   fatty  a.-uls 

l,v    a     fanltv    m..1al...lisni     ^  w    I'M..'.'    IW  ■ .     Tt ss..nt,al    .-aus.'    ..t     th.- 

n-i.l.-sis  is  tln.n.for.'  .'ntir-'ly  .lilT.M-..nt  from  that  in  nr|.lint.s:  ,n  .l.a- 
!„.t,.s  for..i...n  a.'i.ls  are  a.l.L.l  L-  In^  I.I0...I.  wln-r.-as  in  n.-pLr.tis  tin- 
,„.i.ls  of  a  nornial  nu.tal...lism  a.-mmulat..  L.-au  ■  of  faulty  .-s.Mvtn.n 
,l,rou.M.  tin-  ki.l.n.vs.  Tlu-  usual  si-.^.s  ,.f  a.-i.L.sis  oxist  in  l.otl,  .-asos 
l.o,.aus,-  tin-  sur,.lus  ..f  a.-i.l  .l.fl.t.-s  th..  st-.ro  of  l.u.arl...nat..  an.l 
,..,Ms..s  .-lian-.-s  in  tin-  alv.-.lar  CO,,  in  t!..'  ( ■(),-al.s..rl.inir  l...\v.-r  of  II.0 

1,1 1    in  tlu'  r.'s,.rv.'  alkalinity,  an.l  in  \W  a.M.l  .'N.-r..ti..n  l.y  tlu-  Uulncy. 

1,  is  important  to  r.'.-.-^niz.-  tin.  s|....ial  natnr.-  of  .lial...ti.-  a.'nL.s.s  l.y  a 
separat.'  nam.'     h<  texts.  . 

Tlu>  .•h.'n.i.al  i.r...-.ss.'s  l.y  ^^lu.•ll  tin-  k.'t..n,.  1....1i.-s  aro  pr,..ln.M.,l  is 
.liM.usM.I  ..ls..ul,..r..  .pa-,'  7(1!!..  H  n mains  f..r  us  t,.  P..ns,.l,.r  tl,. 
._r,.n.'ral  nature  ..f  tin'  metal...lie  .listurl.an.'o  r.'sp..ns,1.1e  t..r  tlu'.r  ap- 
pearan.-e  in  dialieles. 

Vuv  tl..'  tl,..ronu!.  .•..ml.usti..n  ..f  fat  in  tin'  animal  l....1y  a  .•.•Hani 
,,„„„„„  .,,.  ,,,,l.„l,v,lrate  nn.st  1..-  sin,ultan....usly  l..un...l.     F""'V'-1<;-"  >' 

is  .,  l.'ss  r.'a.lilv  .,xi.li/..'.l  f Islutr  tl.an  su-ar-.  it   n.'.'.ls  tl<o  lire  ..f  tl..' 

,„„.„in.'   su.'art..   -onM.m.'   it.      If   tl,e    ..arl,ol,.v  .Irate    tires   .1..    n._.t    l.urn 
l.risklv  en.,n..'li.  tin'  fat   is  in.'..mpl.'t,'ly  e..nsunM..l:  it  smokes,  as  .1   wer,' 
.n.l  tin-  sm..ke  is  repres.'nt.'.l  in  metal...li-m  l.y  tl,e  k.t.-nes  an.l  .l.rive.l 
,,.i.ls       Suel.    a    el..sin.„'   .l.-NNU    of    tl..'    earl.ol.y. Irate    furnae.'s    n.ay    1..' 
l.n.u.'lit  al...u1   eitk.'r  l.y   eurlaihn.'Ut   ..f  tin'  intake  ..f  earl,..l..v  .Irates,  as 

M,   starvati..n    ^...r   M.\n .   ,.r   l.y   so:,.-    fauh    in   the hantsm    o|    the 

fnrna.'e  itself,  as  in  .lial.etes.     Resides  fat.  J.rotein  .nay  als.,  eontr.l.ute 


'I 


iMii- 


I      !   i 


1.  - 

:* 

Jti 


i;i    \iu'^ 


f    i  i' 


V*  »  I 


684 


MKTAnOI.lSM 


fo  the  ju'ddiictinii  of  kctDiii's  \\ Iicn  ciirlxdiydrnto  ctiiiiliiistion  is  tic- 
jin'sscd.  I*"iiii(liinifiiliilly.  tlicicfurc  kcfosis  in  dinliotrs  is  duo  1o  tli<' 
sjiiiio  cause  iis  ill  stai'\  !itiiiii-  iiiinicly,  iiii  iiiiijnipcr  adjuslmciit  hctwocii 
tlic  iiicfal)olisiiis  of  fat  aisd  carlxihvdrate. 

Hcariiif;  tlicsc  iiiiiiciplcs  in  ?niiid.  it  is  easy  fo  sco  liow  the  intensity 
of  aeidosis  wliicli  dcveltiits  diii'iiifr  starvation  will  dc|)end  upon  tlie  re- 
lative nietaliolisni  of  carlioliydrate.  on  tlie  one  hand,  and  of  fat  and 
l)rotein,  on  the  other;  it  will  therefore  depend  on  tlie  amounts  of  these 
foodstiilTs  which  have  lieen  stored  in  the  orifanisiii,  and  this  ajraiii  will 
depend  on  tlie  nature  of  tlie  diet  previous  to  the  starvation  period.  For 
the  first  few  days  followinj;  entire  alistiiieiiee  from  food  in  a  healtliy. 
well-nouiisheil  individual,  very  few  if  any  ketones  will  he  excreted  in 
the  urine,  hcciiuse  the  carlioliydrate  stored  in  the  liody  as  alyeotren  has 
sufTleed  durinu'  this  time  to  maintain  the  i)roj)er  i)roportioii  between  fat 
and  carlioh\drate.     Aftciwards.  however,  their  aiiiiearance  is  to  he  ex- 

P<'«'tcil,  I aiisc  the  j;l.\  c-o;:eii  stores  hecome  exhausted  loiif;  hefore  those 

of  fat.  If  stai\atioii  is  still  further  inolonjied.  a  stasre  will  come  when 
thn  fat.  iis  well  ;is  the  carludiydratc,  is  used  up  so  that  the  or<faiiism  has 
now  to  sulisist  on  ))rotein  alone.  When  this  staire  arrives,  the  ketones 
\vill  diminish,  for.  althoiijih  they  iiiiu'ht  be  derived  from  certain  of  the 
amino  acids,  yet  this  iloes  not  actually  occur,  hecaiise  a  lar<re  jiart  of  the 
protein  molecule  i  nearly  lialf^  also  hrcomes  ehanwil  into  sfliicose,  which 
liy  liiirniiisr,  as  above  ex))laincd,  prevents  the  formation  of  ketones  from 
the  other  ))ait  of  the  molecule.  For  the  same  reasons,  marked  acidosis 
will  Hot  be  exi)cetcd  to  occur  diiriny;  any  statre  of  starvation  in  1e;in 
persons,  who  from  the  start  iiiiist  utilize  mainly  their  stored  l>roteiii  to 
supply  the  fuel  upon  which  to  liv(>. 

In  diabetes  exactly  the  same  principles  apply,  but  to  ;in  or<;ailism  in 
which  the  iibility  to  metabolize  carbohydrate  has  been  depressed,  so  that 
"the  maximum  rate  at  w  liieh  dextrose  can  be  oxiilizcd  is  fixed  at  some 
level  which  is  absolutely  lower  than  in  health."  "  Therefore,  since  a  cer- 
tain imipoitioiiality  must  exist  between  tin-  rates  of  combii.stion  of  fat 
and  c.irbohyilrate.  the  diabetic  can  thoroiitrhly  oxidi/e  less  fat:  in  other 
words,  an  amount  of  fat  which  could  readily  be  bui-ncd  in  a  healthy  bod\ 
is  imitroperly  burned  by  the  diabetic,  and  ketones  and  their  acids  ac 
cumulate. 

Starvation  Treatment.  "In  order  to  check  a  di.ibetic  acidosis,  it  i^ 
necessary  to  restore  the  proper  ratio  of  fatty  acid  to  <rlucose  oxidation." 
which  e;in  best  be  done  by  starvjition,  rest  in  bed  and  warmth.  Hut  this 
treatmeiil  m;iy  not  at  first  suflicc.  because  we  have  to  deal  not  only  with 
the  acidosis  bodies  deriv.'d  from  fat.  but  w  ith  those  which  can  be  derived 
from  protein  on  account  of  the  diabetic  ort;aiiism  haviiig  lost  the  powei 


Till.    MKTAIIOIISM    OK   Till:   CAKIIOIIYDRATKS 


fiSf) 


ovon  of  LnrniM!:  tl...  «lu.-.s..  xvl,i.-h  is  .l.-riv.-.l  I'n.n.  Il.is  t.M,.lst..tT.     1  y 
,K.,.sis..Mu...  in  tlK.  starvation,  howovn-,  tl..  al.ili.v   of  th.  o,.«an.sn.   to 
;„ili...  ..arLol.v.lrat..  nsnaiiy  l,...-on,..s  so  far  .vstor..,!  tl.at  .■nou-.h  Imrns  to 

,„,.,,.,„  ..n.l..is.     Kv..ry  .as.,  of  .lialu.t.-s  .-an  not.  tlx-rHor...    - '"''l--  <■; 
„  n-a-t  in  tlu-  sa.n..  way  to  starvation,  tl,.  .iHornmun.  .■on.l.tn.n  Wu.' 
,,^^.  ,.„,^,,i.,„  ,„,„..„.„  „,..  ,,„„„i,i..s  of  «ly.o...n  an.l  tat  stor.-.l  m  tin-  l.o.U 
a,  ,1...  onts..t  ..f  tin-  fasting  l-rio,!.     This  r.-lationsln,.  <l.I..n.ls  on  tl,. 
natuiv  of  til.  pr.'vious  diet.  ,      ,,         •      i-  i    *.. 

To  sum  up,  "fasti...  will  low.r  a.i-losis  .i,l..r  i..  lusiltl.  or  .n  .l.aL.t  s. 
if  it  has  th.  .tT...t  of  stoppi,,,^  a  on.-si,!..!  .n..tal.or.s.n  an.l  th.-..w,nir  th. 
,issu.s  ..n  a  ........  n.arly  l.ala.....l  rati...  of  fatty  a.i.ls  au.l  du-os.-    -^- 

(Wo...lvatt).    A  P.'a.ti..al  p..i..t  n.ay  h.  not..!  h.r.- ..an..- y,  that  t  ..■ 
s  lik.lV  to  h.  ...O.V  .la..v'.r  of  MM^inus  a.i.los.s  .l..v..lop.n,'  .lur.ni.  st..,v..- 

,;.,„  i,;,-,,  „,nn  i.i  l.a.,  .lial...ti.s.    Tl,.  in.po.-tau.o  ..f  .mr  apprca  lo..  ot 
,U.s.  fa.ts  in  th.  starvation  t...at.n.nt  of  .liah.t.s  w.ll  h.  s.ll-ov,.l.ut. 


hi 


■. 


i 


1    I         Ji 


'l;li  '  U 


■  :    t 

it: 


ih 


i  : 


iM' 


iil! 


liiMI 


!  i'  r 


H     ^ihVf 


U' 


'ii 


m 


i 


(  II.M'TKi;   I.X.WII 

FAT  MKTAI'.OI.ISM 

Hi'fiifc  coii-iilfrini:  till'  i>livsiiili)uy  of  f;ils.  ;\  few  of  tlic  nmst  csscntiiil 
points  rctrfiiiliii'_-  tlicir  cliciiiiNtiy  may  lie  of  jissi>t;iiict'. 

THE  CHEMISTRY  OF  PATTY  SUBSTANCES 

It    is   llslllll    til  I'lilssil'v  .-ill    MlllsliiniTS    lllilt    Jill-  Millllllc    ill    CtlllT   ilS   ll'lltlills. 

They  iiirliulf  |';ittv  jiriils.  iinitiiil  f.ils.  rlinlfslciiils.  cliiil.'sti  rill  I'sti'i-s,  ami 
l)lins|)Iiiili|)iMs. 

Till  hilhi  mills  lii'liiip.'  til  t\Mi  iiiiiin  liiiiiiii!iiiriiii>  sciir^.  w  hii-|i  ilitTrr  I'lmii 
(Mi-li  iitliiT  with  icMMiil  til  wlii'tliiT  tlii'v  ail'  satuiiitril  iir  iiiisatiirati'il.  A 
siiliinitiil  fatty  ariil  is  typilinl  liy  palniitii-.  wlinsr  rnriiinla  is  < 'IF  ("IF  .-('I!  - 
<  IF(FF_  (FF  (11.  CM.  (11  .(  li.CJF  (11,  <II.-(M  -CII  (( )( IFF.  or"  CIf" 
I  ("IF  \  ,-( 'OOIF  :  that  is  to  say  it  is  a  lii^'ln'i'  iiiiinliM'  of  tin-  si-rii's  to  wliirli 
ai'ftii-  ai'iil  iCFF  CdOll  >  Ih'Iomu-s.  (jilliriii;.'  from  tlif  iatti'r  in  having'  fmir 
ti'iii  I'Ntra  nictliyl  laiiii-li's.  rarli  joiiiiMl  to  its  nci^'lilior  liy  niic  ImhhI  i>i'  satii 
ratiil  linking'  mi  citlirr  siilr.  Aiiotln-r  nimiilicr  of  tliis  scriis  is  sti'aiii-,  in 
wliii'h  tliiri' an' sixtiin  i'\tra  CII,  •.'fmiiis  i  CFF  ,M 'IF  , ',,('(»( »FF  1.  An  im 
siiliiniliil  fatty  ai-iil  isi.j.i,-  iCIFM'IFi.  CFF''  (  IF-  ( 'IF  .K-( 'OOIF  ^. 
Its  iiiisatiiialiiin  is  iiinismiti'il  in  tlir  furiniila  liy  tlii'  iloiililc  lioiiil  or 
iiiisatiirati'il  linkiii'_'.  wliirli  it  will  l.i"  m'i'ii  oi-i-npii's  a  position  in  tlio  niiiF- 
illi'  III'  till"  mnlii-iiii'.  till'  otiii'i'  mctliyl   raiFirlcs  lii-iiiir   linked   tofji'tln'r  I'X 

sllliilc    lllllllls. 

Till-     fatt,\-    ai-iils     ri'ailiiy     i-nmliiin'    with    alkali    to    rorni   S'xf/i.v;  llilis, 
CII  (('II,i,,('(M»II      Kdll      (11    (11.   ,,-('()(»K      ll.(t.   till'   naition   iM'iii- 

f  palmitii'  ariil  i  (soai)') 

analoiroiis  to  that  liy  wliii-h  ai'i'tli-  ai-iiF  forms  an  ai-i'tatr  with  alkalii's. 
Fn  plai-i'  iif  liiiiii.'  I'liiiiliiiii'il  with  alkali,  tlir  ('(KlFF  I  I'arlinxyl )  <:riiup  of 
fatty  aciiN  may  rumliini'  with  alrohols  to  furiii  siihstanccs  calli'iF  ixlirs. 
Tims.  ;i"i'ti,-  ;i,-i(l  ■iml  itli.\l  ali-uhiil  form  i'tli\l  ai'i'tati', 
IIK'IK.  II  oil  (',11  ('IFr(KM',IF  Flo.  When  tlu'  alcoliol  lliiis 
'"'•I  til-        ^lihy!  'othyl  acftate) 

ariil  1         jili'oholl 
uiiitcil  with   fatt,\-  ai-iil  is  >rl\iM'inl  (jilyci'iiiii'i.  in  wliii-h  tln-rc  arc  iFirco 


KAT    Ml  TMKil.l-M 


r.M7 


(.11    ll,,v,lr..sv|.    U'lnnpS    )ll.'    ITMlHin..'    rslrr      .illl..!    tliL'lvr.li.lc^      is    »M». 
/)•.(/  /"-//.     'rripalmitiii  luis  tin-  I'l.iiiuilii : 

(11      (>()<■•••  II  , 

I 

(11       (MT  (•   II  , 

I 
(11     ()(>(•(•  II  . 

r.v  iM.ilil,..'  liH.tr;.!  l-.ts  Nvith  ;,lki.li  tl,c  r.tl.v  ;,c.l  iM.liMcs  i.n-  H'l'f  "l^ 
..,s  sonps.  Icavinir  tlu-  «lyn-n.l.  This  ,.nM...s.  is  ..nll.-l  M,,.„M/...;/y(.-.  an.l 
;,  ,„,,v  1h.  ..IT..H...1  in  .,.,.,>  „llH.r  u.-ivs,  ns  f..r  cxa-M,.!-  l-y  lu.aln..'  xvitl. 
st.M.n  (u-  l.y  tli-  a-ti.m  ..f  S'-i^'l  "">'> '-"-  '•^'"'"1  '''"'"  ^^  ^^''"■''  '"'•'  '""''• 
(listriliiitrd  in  plants  and  animals. 

.|-|„.  natural  fats  arc  usuallv  a  niixturr  nf  iri..'lyccri.l.s.  and  their  -lif- 
,V,.,nc..s  in  pnMMTli.'s  arc  .l.p.n.l...,!  np.n,  tl,c  r-laliv,.  ann-unts  ,.t  fatty 
,,..,  Is  present.  Tin-  three  in-st  important  in  animal  lats  are  tnpalmit.n, 
tristearin  an.l  triolein.  It  is  essential  in  the  study  of  fat  metahohsm  that 
we  should  know  the  most  in,porta.,t  n,.lhn,ls  h„  irl.irl.  Ihr  ,.n.,><>rlu,„  nl 
hlhj  n.i,h  ,.,:.r„t  In  a  mn.l  ht  />•  <hlnwnn,1.  Those  in.^tlu.ds  are  as 
follows: 

1  '/■/,(  m.irnni  ,„-n,l.  Oh.in  is  li,,nid  at  «  C. :  palmitic  aeid  melts  at 
r,-'(;  (•  •  and  stearic  at  (iK.:!  C  The  solidity  of  animal  fats  depends  on 
,1„.  |,rop,.rtion  of  olein.  i^almitin  and  stearin  present.     Mutt.u.  fat.  for  cx- 

;„nplc.  is  much  stitVer  than  ).i-  fat  heeausc  it tains  less  olen.  and  nn.rc 

stearin.  The  meltin-  poi.ds  of  fats  fnmi  ditTcrent  parts  of  the  hody  nn.y 
also  varv. 

o    '/•/,(  „rh1  ,„nnhr  indicates  tl nmnnt  of  free  fatty  acid  mixed  xvith 

the  fat  and  is  determined  l.y  titratin-  a  solnti.m  of  a  Nvciu'hcd  .p'antity  ot 
,l,e  fat  in  alcohol  with  a  N  Id  alcoholic  solution  of  KOII.  phenolphtha- 
lein  hein<r  Used  as  indicator. 

n    Th,    s.i,,o,nli.:,ll<.,i    n,h,.    indicates  the   total   amount     of     fatty     acid 

present    l-oth  that  uhid,  is  free  ami  that   e.m.hin.'d   with  triycerol.     It   is 

determined  hv   heatin-  a  weighed   amount   of   fat    with   an   exactly   known 

i„Mon.>t  of  alc.holie  KOll    i.h'termimM  hy   titration   with   standard   acid   . 

\fter  saponiticati.m  is  e-uni-lcte.  titration  of  the  mixture  shows  how  much 

alkali  has  1 n  used  t.,  e.md.ine  with   the   falty  acid.     This   is  the  sa|.oni- 

fieatioii  value. 

4  The  rsl,r  nilin  indicates  ihe  amount  of  falty  acid  c.mil.ined  with 
>rlyeer(.l,  ami  is  ol.tained  hy  suhtra.tin-  the  acid  value  from  the  saponi- 
fication value. 

Ucsides  these  there  are  luw  values,  kno^^n  :,s  llie  iedine  and  tl..'  Keicl.ert 
Meissl  values,  that  are  of  importance  l.ecause  th.y  depend  on  r<rl,in,  ,-ln,r- 
iichristlnt  of  thr  faltji  ncii!  jwllilis. 


I   H  4 


y 


!Hn, 


itii 

•  Hi 


hi     li 


hi 


.1- 


1  I 


(ISM 


Ml  TMMil.ISM 


ilii 


!i    t 


!>■'   H 


,").  Til'  linliii,  nihil  imliiMlo  \\i<-  iiiiioiiiit  of  iinsiitiiriil«'<l  t'iittv  m-iil-i  |>rrs. 
.'lit.  (ir  till'  iiiiiiiImt  (.r  cldiil.l.'  I  i.ihIs.  It  ilr|.i>ii.|s  (III  tin-  I'ii.-t  that  i>.iliii.', 
like  iiiaiiy  (itliiT  Milistaii.f^,  is  cai.iil.l.'  ..I'  .liivcilv  attarlmij.'  itsril'  I..  ,m.' 
falt.\  arid  i-hiiiii   \v1ic|(\ct  ilimlilc  1  ninls  csist, 

(1.  77m  ll.iiinil  Miixsl  nihil  iiiili.-ates  tin-  aiiuMiiit  if  volatile  vlilldr 
nri-l  piTscnt  ill  tlic  lal  It  i.  .l.'tn  iiiiii.Ml  \,\  liisl  ,,t'  all  >a|>oii'f\  iiitr  lli'- 
fat.  tlicii  (lcci.iii|iM-iii'.'  till'  >«ia)t  liv  niixiii'.'  it  uilli  iniiinal  a.  id  aii<i  "lis 
tilliiii:  the  lilu'i-at.'d  l'att.\  ari.l.  tin-  .ii>tillatf  luiii'/  ci.ll.'.'t.'.!  in  a  Uiiovmi 
aiiic'iiiit  of  stall. lanl  alkali  an.!  tittalc.l.     It  is  a  valm-  tl.it  is  ii..t  ..t'  \.r> 

L'l'.'al    lis.'  ill   pli.v  si.ili.L'ic   iiivcstiuati.iiis,  luit    it    is  s.i  ii lui.'.'ti.iii   willi 

f 1  cli.'iiiistn  .     Sill.'.'  volatile  aei.is  are  |.!('s.iit   in  liiltter,  tile  K.'ii-lieil 

M.'issI  valiii'  li«'l|>s  lis  t"  .listiii«itisli  ln'tw.'.'ii  Imtti-r  and  iiiar'.'aiiiie. 

Kat  is  iiisoliiM.'  in  «aler  l.iit  s..a|>  is  s.iliilile.  f..riiiiiiL'  i Ilni.lal  s..lil 

ti.m  wliieli  ]ires.  iits  till'  pli.'iioiiK'ii.in  .if  Mirfai'e  ayufeuatimi  id"  iiii>leeule<. 
This  .'.insists  ill  tli.'  .■..iieeiitiati.pii  "f  tlie  s..ap  imtli  at  tlie  free  siirfaee  .if 
tlie  li(|ili.l.  wli.'ie  a  skin  may  fonii,  aii.l  at  the  iiiterfaees  lietweeii  tin- 
s.iap  s.iliiti.iii  aii.l  aii.v  liii.iiss..|ve.l  paftieles  ])i'eseii1  in  it  This  pelliele- 
f.U'iiiati.m  aniiml  thi'  |)aitii'l<'s  prevents  them  t'nnii  nimiiiii,'  1i>._'eth(M-  s.. 
that  tli.\  r.'iii.iin  siisp.'ii.h'd.  thus  iViriiiiiiir  an  i  iniilsinii.  .\ii  eiiiiilsi.ni 
may   tlii'rel'.ii.'  In'   t'liiiiie.l  I'ither  til"  n.'iilral   fat   of  any  otlnT  )ihysieall.\ 

similar  sulistai Wli-'ii  fat   itself  is  iise.l.  there  is  usually  eiiousrh  f i .'' 

fatty  a. -ill  adiiiixed  with  it  l.t  iiiaki'  it  uiinee.'ssary  in  t'ormiiit;  the  I'liiiil 
sioii  t.i  il.i  III. .re  than  shak.'  the  fat  ui'h  w.'ak  soilimii-earlioiiate  solution 
With  .ith.'r  sulistaiiees  ii.it  eolitainini:  any  fr.'e  fatty  ai-ni,  somi'  soajw 
slioiil.l  h"  a.hle.l.  To  pri'sei-M'  the  emulsion  it  is  .(fteii  useful  l.i  add  soiin' 
mileila'-'e.  In  th.'  eiiiiilsilie.l  stat.',  ii.'iitral  fats  are  iiiiieh  more  reailily 
atta.'keij  hy  lipases  than  when  they  are  |)'esent  in  an  iiiieiiiulsilied  stall' 
Thus,  emiilsiti.  .1  fats  an'  ".ru.'este.|"  liy  the  lelativ.'ly  small  anioiiiits  of 
lipas.'  present  in  the  stoiiiaih.  wh.'ieas  neutral  fats  thenis.lves  are  not  so 

h'alty  aelils  .ilso  .'sist  In  iiatur.'  in  .•oiiil>inati"ii  not  with  the  triat.iinii- 
aleohol.  <^'ly.'.'r.il.  Init  with  monaloiiii.'  ah-ohols  siieh  as  i  huh  sli  ml.  Tliesi 
eli.il. 'Sterol  fats  .lilT.r  from  the  l'I.v  eer.il  fats  in  heiii.;  v.-ry  resistant  t.. 
wards  eii/\iiii's  and  miernoii.'aiiisiiis.  They  a:-e  therefore  used  f.u'  \>'>i 
te.'tive  jiiirposes  in  the  animal  eeonmny  ;  for  example,  tiny  oeeur  in  the 
selium.  the  seer.'ti.iii  ..f  the  s.'liaeeous  ..'lands,  \\heri'  they  serve  to  inoistili 
the  hairs  and  skin.  They  are  also  pres.'iit  in  eells.  in  which  it  is  proh 
ntile  they  tiike  an  iiiiportaiit  part  in  formiiii;  the  skeleton  of  the  cell 
Cliolesterol  is  alisoilie.l  from  the  intestine:  it  is  always  present  in  the  Moo.l 
hoth  in  plasma  and  in  eorpnsi-l.'s ;  and  it  is  an  important  eonstitiieiit  of  hile. 
from  wiii.-h  it  may  s.'paiale  ..lit  in  tlie  hiie  piis>ai:es  and  form  eaicul: 
((lallstolies). 

Ill  the  eells  theins.'lves  the  lipoids  are  repri'sented  iiiniiily  hy  poin|M)uiid- 


fl 


11 


iiu^ 


y\Y    Ml   IMUil  l-M 


f.80 


,.!■   :,  s.„n.'ul,.,t    „,..- ..|.l--v   v..,..l..r..      .w. I.v,    th-    ,,l..,.,,hnh,„„..      Ah 

,1,,,,,  „„„..    ilHii.,.!-.  Ih. i^iM   .  t.-Mii.allv  ..f   |.|M.M«l".nr  .ir,.l  ,M.ml..ll-.l 

«itl,  H.MiIn.l  hiX  nn.l  «nli  m  iHtn.ur..„.,u>  I  .i-'.  .l-li".     'H  -  ""•>'   ><"■'""'  "' 
,1,.,  ,.|,...i'li"lM'i"s  iW,-.f/.M,.  wlu.-l.  is  wi.lrh  ,lMr,l.ut-..l  .„  th..  i.i.Hnal  I^hIv 

,,,..s,.nl    in   1.1 1   i.'Hl   1m!..  ii-  u.  II   as   It.  all   .•■■lU   .     OiImm-   ,,t,..s,.li..l,,.ii.s 

,„.,.^,.>.l   in  n.iv..,,.  tisM,.    a..-  -..i,!,..!!.!.  .•imrin  u.hI  >,.hinu'-tnv  .li.i      TImt.- 

„,,    variciis    l.v.ilniis   .l.stin..'nis|„.l    f- ..MotlHT    l,>    tl.-    latl.v  a-i.l 

,,.li,,|.s    NvlM.li    Il;.7 Mi"        l>,~l-ar>ll.-ill,M,.    l-r    .Aalnpl...    lias    tli.' 

I'..'  inula: 

I'M      I)    (II    ril     ,    <'ll, 

t 
ni     o    (><•  I'll        I'll 

I  «li  :iiii-  liii'l  I 
(•II,      I)  () 

'        \     / 
f-lv,.'n.|.        !• 
/    N 
nil  11'  II      <'ll       "^   '"  ' 

.  |.|i...|.li..ih' 

,„i.|)  '>" 

..'hul'IK-l 

This  ..,„„|,l.'s   ninl.'-ul.'  .-an   na.lilv  1,..  .plH     M-  ''>    l'V'l''"l>si-^   Mvannin- 
with  liaryta  wal.T'   inln: 
-Iv-.-iM  |.!„,-|.I,..M.-   a.i'l.  ''11       <•" 

(11  -  on 


ni    o        o 

\  / 

I' 

/  \ 

oil  OH 

l,,,|i,,\i,l..  1  ;    nil. I    f:itl,v    ;i.i.N. 


;.l,..l.ii.'.  N 


(r  ll.oll 

I'll 
nil 


itlivl  aiiiii.Hiiiiiiii 


Willi  l,^.ll'u■■l,^.,■i,.  aria,  -Lolin-  f.-inis  a  salt  ul.i-li  n'a.lily  f-.iM.s  n 
,,„„1,1..  salt  wi.li  l-tatiiii-  -lil-.i.!...  Sin,.,,  tl.is  -l-ul.!..  salt  f,.nns  .-lian,.- 
t,.,.is,i,.  ...'vstals.  it  is  ns..,l  t,.  i,l....tilV  an,l  s..,.a,'at..  U.-itluns.  !■  „r  .p.an- 
,i,„iv..  ,;,u-,ns..s.  l,.,u,.v..|'.  it  is  ninn.  si.ital.l..  I.,  a-tcnnuu-  '-.M.l.n,  .n- 
,li,,,.tl.v  hy  fill,  aiiinunt  „!•  ,.h..si.i.„vi..  .i-i,!  I-— '"t  ">  ""  <'""''''"'  ''^• 
tia,.t  i>f  IIk'  ,ir'-'aii  <>r  tissii,. 

Kvi.l,., is  ....nsta.itlv   a.-..n,ii>latiiiu'   t..   slnnv   that    li',-ithin   is  an   ox- 

.n.nn.lv  i tant stitu.'nt  ..f  ,-..lls-.  iinl 1.  it  s..'    s  tn  ho  th.  intof- 

,„,,Iiat",.  sta.-M.  in  th,'  ,il  ili/ati-n  of  iionliat   fat.  hy  ,.i   .■„.|.las,«.     Its  ,>hns. 
,,l,„,„s  als,.  ,.i'.,hal,ly  s..,'v..s  as  ll,-  s.ain...  of  th^-   ..|..n...nt    f-,..  tho  .on. 

stru.-.i r  nn.-'..i,-  a,-i.l    i  s,'.'  V.^'^"  <-T..      In    airvus  tissu-s   ,t    is  ott.n 

.....ial.-a    uill:    ^.arh^^hy.h'Mt..    ,no!..-Ml.'s    ( >.ala.-tos.. -.    foniMn^   th.    s.ih- 

ici'd'orc  liavo  soiiii'  I    If  to  iilay  i" 
I'kiis   also  attrilniti'   to   lecithin   an 


NtaiKM'  known  as  ccfchiiii 


It   ina\    tlii-rcl,'!'!' 


ilM.liv.liat,.  mclalMilisiii.     S,iiiH. 


iP  li 


i'  !' 
1 


i  1 


t 


it 
I  I 


I-I' 


f  i 


Hl!i 


it!     '  i 

\r    1 


I 


1 1 '       1 
IS 


I 


1,^ 


;    ■    ( 

I    ; 

'   I       ! 
1 

*        ! 


■•'.I 


M 


\\h 


tft.i  I 


t;!Mi 


Ml   IMlllI  l»\I 


iM,|.nll;.Mt    I'llliHinn    ill    ill.'   1 1  lUislrmHr   ,>(   Mll.-h,  IMMs   lliluuuli    r.ll    MMin 

l„;iiH>      WIm-ii  iiiiM'.l  ^^  iMi  uiil.f  it  su.lU  u|.  I'V  iniliil'iliuM,  mikI  it"  nv- 

lallniiN  Of  nlli.r  su^■^l:ll »  ;ilv  .|i>N,.l\  r.l  ill  tlir  Wiitrr.  :i  iii.'Jills  is  olT.T.'l 

f,,|.  iMinviMiT  „,il.i- -i.liilil''  iiH'l   fill  N'lliilil.'  Mil.xliiiir.'s  into  iiilinml 

Illrf 

DIGESTION  OF  FATS 

A   r.il.iiii   .•HIM. lint   ..r   l':it.  .siM.'ially   u  lun    it    is  in   nil   ri„iil.ili.Ml   .•..ii.li 

tinh,    ,MI1    I...    ,|;-rslr,|    ill    111,'    sl.MILI.'ll    I'.V     IIm'    li|'ils itaill.'.l    ill    til.'    U'il- 

Mi,.  jni.T       Mnst    ..t'   it.   Imu.M  r,    i.  ,li.jr>t,.,l    1m   Hi.'   mmmII    iMt.'stin.'.    int.. 

^^lli.•ll  a-  v.'],:\\<-  m-h.  H   i-  '^i  a.lu.'illv    .ji^.-liai -.  d  siisi.,  li.i.'.l  ill  tl Il.\ 

\\,y   tlii>   liil.'stilial  .lii;.-^ti..li   '<(   lal    Imlli   i„i  iirr<  <ll 'f   Ji  in    "inl   I'ili    iiri    in  r 
,.s, (,-./.      '11. !>   l'^  .mmIv    sIiuum    ill   til'   ralil.it.    in    vvlii.-li    tin-   |.iiii.T.'atii-   .Iil'l 

iiitirN  III.-  iiil.-.liin'  at  •■ iisiiliialili'  .l^tai I.cl"."    tli.'  I.ilc  "lu.-l.      If  'li.' 

111. ■Milt. ■!■>    is   in.|v.-l..|    .jiiiiii.j    III.'   al.s..i|.ti..ii    ..f    fatl\     \'>"<'\.   M"    I'"!    i" 
i,.,'li..ii   ..f  til.'   I.Mii|.liali.-.   uill    I"'    ii.'l.->l    l..'t\M'.-ii   111.'   I'll.'   an. I    tli.'   |.an 
.-(•.■atir  .lii.t^  I. Ill  ..iil.v  !..•!. .w  til,'  l.itt.T,     III  til..  .I..ir,  in  wlii.-li  ImiIIi  tli..  I>il.' 
an. I  111.'  main  |.aii.-ii  at  I.'  .Iii.'ts  .'iil.T  tlic  int.'Nliiif  at  ill... ill   tin'  >-iiiiic  l.'\.l. 
fill    iii.i....ti..ii  .,!    111.'  I>lll|.llali.••^  stalls  at   tliis  i...int.  I. Ill    il"  tli.'  Lilt'  <ln.'t 

M.r  latli.T  111.'   uall    l.ia.l.l.r^    is   Iran   plant. ■.!   at   s listi I..u  n    tin- 

iiil.-^liii.'.  il    will   I"'   f..uii.|  lliat    111.'  inject  i. .11  ..I'  till'  Iviiipliatii's  willi    fat 
CI... •Ills  i.iil.v   l..l..\'    ill.'  11.  \\    |'..iiit  ..)'  iiisiTti.in  lit'  III.,  liili'  'Iil.-t. 

K.'tii.ival  i.r  II!.'  paii.T.'a^  int.'rf.  r.'s  v.'i'v   inaL'tially  witli  tli.'  alis.ir|.ti..n 

,,(■   T;,!.      In    Mian.    t'..r  .•Naiii|.l.'.   al.s.'ii. f   tin'   pan.  r.'atic   .jni.-.-   al.in.>   ili 

iiiinisliis  111.'  ali^i.ipti.>n  fat    l.v   .Mt  <>v  (ill  p.T  .•.'lit.      If  tin-  Ml.'   is  iiK". 

al.Miit,  111.'  .linrniiiti..n  aiii.iiints  t..  HI  ,,r  !MI  p.  r  .•int.  .'iii.l  in  '-n.'li  .•ascs. 
a>  is  \wll  kii.nMi.  111.'  a.liiiiiiistfiili.in  of  Ml.-  .>r  pam-i.'.'is  powil.T  '.'ifatl.N 
iiiipi..\i-.  fat  al.M.rpli.-n      In  tin'  <|..i.'.  altlnni.-'li  li'-Mti..ii  of  tin-  pan.'nali.' 
.111. 'I    appai.Mtl.v    ..iil\    sKn^ilitly    inlln.'ii.'.'s    fat    al.soipti.in.   v.iii.i\al   of  lli'' 
liaii.'i.as  iNi'lf  u'f.'all.v    int.'it'.'r.'N  with  III.'  pro.'.'ss;   fr.mi  wlii.-li   fact   soni.' 
ol.s.'iM'is  li,i\.-  .•on.'liKJ.'.l   tliat    111.'   pain-r.'as.  in  a.l.liti.ni   to  its  cNti'inal 
si'iT.li..ii   into  111.'  int. 'still.',  must    pro.ln.-.'  an  iiit.'tnal  si'd'ction  into  tin 
lil.io.l    \\lii.|i    lias   soiin'tliiiiir   to   '1.1    "itii    til.'   .'tli.'ii'iit    alis.ii'pf ion    of    lli.' 
fat     rralt,  M.I  Inn   an. I  \'ln.'.'nt '         Il  is.  Ii..ui.vit.  inippilial.'.'  tlial  sii.^li  an 
li\  p.itli.'sis  is  ii.'.'.^ssary.   f.>r  it    is   very   lik.'ly  tliat    tlif   niorilnin.l   con.li 
tion    into  ulii.'li   an   animal   is   liron..'lit   liy   I'xtiipation   of  tli.'   i.ancrcas, 
a.l.-.pial.'ly  a.'.'..unts    for  tin'  siippr.ssioii    of  tlic    fat  alisoiliinu;    fiinclioii 
.\s  t.i  llii    riliilii-i    roll  ft  iif  iiiiihrnitir  jiiin    ami  hih    in  tlic  .li'.'cstion  of 
i'al.  «.•  know  of  .•oiirsc  tliat  in  tli.'  paii.-i'.'iiti.'  .jiii.-.'  tli.-r.'  exists  a  lipolyti.' 
cii/.\iin'.   liii'i^i.    uiil.-li.    iiii.t.'i-   siiilal.i.'   c.pn.lilions   li.i.s   tin-    p..\v.  r   ct    s-[>lit 
tin./  iii'iitial   fat   into  fatt.v    a.^i.ls  ami  ..'lyccriii...     If  l.ilc  is  cxaniinc.l.  no 
lip.ihti.-  cii/vnn'  will  lie   foiiml  in   it.      It    is  entirely  inactive  on   fat,  Imt 


r  \r  Ml  TMU'i  i-M 
,,,,.   H.islul,.   urn,    /-.-/.    ,,:uu.n...i.■iul...^vMM,    l.vi.-lf..Mlyl-l> 

,.;i,.....,ivi.v    .,nl„.   ,'^ ,..,.,... ini,.,.,      1.    I..^l n    ,„,.,„|.lK„    U,..v.lN 

„,•    ..1,„!,,1,-  ':h.,.1.    II ..I1..1    Mr    salN.    .-.-    -1-    ■ ^"""'.ts    o)    l.,l.. 


riillir-<    ■'I 

t|i;il     llilc    iliMi'il    to    |.illM'l'r' 

Ion   i\  iilr.il    iicliv  aliii'^ 


.,„.,   I.       ,  111..  Il.lltlill  lilt  tll.-lt  llil- 


„,.,    .„...■.•>, Ml,l..    ,„,.,l,i,   :H..n.,.l.n    .,,•   Ii,.as,..   tl„.    f.-.     -vi.iu   1 n 

.1..,, ,t,at..,|«i.ll   l,il-s,.,ll.   I't'lM...!    in-..-lia    u  a>    tlut    t  Ium  -   .  a.  .m. 

;„„„„,„,      „    i,   i,n,H.i.„M    ,„,..|,M,nl...r.   1.-,......   tliat    li,.,-i-..-    it-  M"' 

;,,,.,.  tl,at    lias  M I    fur  ^<'W  tmi''.  liii^  » 

,l,:in    Ml.'    aiiil'MJ    to    fiv^ii    .iui<'iv      It    i^ 
,„..,l,al.i..tl,a..li,-a..t,  n-         '  ■-,.  .,!..  is  .Iik  to  so,,,.-  pl.yM- 

rli.'liliral    rliniiu-'    'umI"         "■    ''    -  ■>''"'■■  .       .    . 

,.    ,       ,  .      ,,...,      ,,    ,.  ,  'r     HmI    |.aiMM.>alir   .1 lliv 

( ».H'  .iia.\   ask   no  ■  ...    •>  ii     m   ■         '  < 

lioth  al'si'lit   r.-oii     '  I    -'   '•       '' 

iiiMlial    I'al   liiit    1  '    '  '  '  ,  ,       .1      •    .      4;.,,.l 

,    ,;       ,;  ,1 '  ,      ...li,,    ,   .■sa.'l.'.l  on  I,;,   tli.'  i.itrstuial 

isfaliiil  il.V'isti.  ll>"     ,  I        s  1  ,1  I-.-    . 

..a.t'na.    partn     '    ■.     ^      : -      ''■-'•'-•      '•'"'■■'•    ''V, T 

,,ou..w.|-.    tlH.    l-at..       .M      I.'      ■      Vl-    ..■V     snn,    al,so,l„  ,1.    I alls,    t  i. 

.,,ill„.|i nh-l •.,;,-..,  ,'.       ..    n„alt,a.l-Mnnot,.,.|to,.,„tl,is 

luni'tioii. 

U..si,l,.s  assisiin.   tlH.  a.  •    ^r       •  ''il"   'V'lit^'t.s   lat   .liir.^stmn   -n 

„,,,.,,    „,,.       Tln.s.    I,^     ,1s    .-ontaininu'   alkali    an,l    nii.-in  liU,.    suKstain-... 
i,   asslsis"in  tlM.  ,.,niilsili,-alioii  .„'  ,a1,      Altlionul,  ..niiiKili-Mt  -n    i^  n-  .- 

s,.nlial    part    of    fat    al.so,,,,  ion.    vH    it    .nally    fa.-ilitat.s   tl,..    ,..■ s.   l.y 

l„...aMn..'    up    tl,..    fat    into    s,„all    .Jnl.nl.s    „i,    «l,iM,    tl,..    hpas.   -an    ..-t 

.„„.-l.    n.o,.,.   ,.ni,-i.„.ly.      Tin.    alkali    aU n.ln.l-s    .itii    tl„.    fat,.     a.-,,R 

as    tln.v    a„.    liK-iat,..!    I..     tl„.    ,li...s,i.,.    pr ss.    ,o    I;""';;'";';"'"';" 

soap..ul,n-li...--a'li'y   :il.^..,.l„.,ll,>    ,1„.  .pitMial   -.lis        I  h-   Inl-  salts 

fi,Hl,-r  assist   in  tl,-s„|,.,i ftl,-falt>    a-i.U,  a,nMl,->    lou.v  tk-  sur 

(.„.,.  ,,.nsion  of  llni.ls  in   vvl,i-l,   ,l,->    a,-  -on.ain-l   ai,.l  so  l.r,,,..   tl,-   tat 
aii'l   lipas-  i,,to  rlos-,.  -i.nta-t. 

ABSORPTION  OF  FATS 

\f,-r  its  ,ii.^-s,inn  tat  li..s  in  .-onta-t  uilt,  tin  M-stinal  l.or.l..|-  ..f  tin' 
,,,,i,l,..|ial  ....lis  as  fatty  a-al  a„.l  uK.-nn-.  Tl,.  ally  a-i,l  is  -omUin..,! 
.i,l,-|.  vith  alkali  to  form  n  ^^  at..|--sol„M-  s.ap.  or  uiti.  l.il.'  Nalts  to 
,•.,,,„  ,  ..,„n,,o,„nl.  xvl,i-li  i>  .-.Iso  solul.l...  Tl,..  .Jy-..ri.,.'  ii.nl  tin'  .lisso!v-.l 
fattv  a-i.ls  ar-  s-paial-lv  aUso.-l.,'.!  into  tl,-  -pitln-l.al  .--Us  ot  tin-  m- 
t-s.ii,-.   in   tin'   protoplasm   of   uhi-l,      aft-r   ,1,-    fatly   M-i,l   lias  l,-.'n   s.-t 

,.,,,,.    ,•,,„„   ,1,..  alkali  en.   I.il-  salt      th-.v    Im ni-   umt.'.i   or    , -s.M,ll,-s./-.t 

,„  ,•„,,„  ,„.utral   fat.  ^^Wu■U  u'.a.lually   lin.ls  its  uay   l,y  tin'   .•-'•ilfal    la.- 


Hi 


I 


■.!i9 


'  ''! 


1:1 


fifCi 


Ml   IMuil  l>M 


Hi 


liiMli 


i>  \i 


li 


V      i   1    I'  ■ 


I      M', 


It  :|   i'  t' 


81    H 


i'      t  '' 


ti'jiN  iiilii  111"  villi,  .-md  llicH  li>    wiiy  ut'  lh4'  1,\  iii|)liatics  to  llic  lliocacii- 

•  llK't. 

TIm'  ilii'iiiii.il  i'\|>liiii,iti(iii  111'  tlic  iili^cip|ili<iii  III'  i':it  is  vii-v  •lilTi'iiMit  froiii 
that  t'co-iiiii-l.\  lull!  liy  liistiil(iL'i>ts  wlci  iii:iiiit:iiHi'il  that  the  line  particles  id' 
fiiiiiUili'il  t'al  ill  tlic  iiili'stini'  |iiii.lrali'  li>   a  i liaiiii-a!  in ess  tliioiiirli 

llie  slii.iti.j  iM.r.ler  nf  the  .•liitllelial  e.'lj  illld  ils  pn  ltii|ilaslll.  Tiie  llistlllilirie 
eviileliee  t'nr  tliis  \  iiw  sciijiei)  \eiy  <'ii|l\  ilieiri'/.  I'd!'  lilM"  fat  !.'liilillles  (Mil 
leaililv    lie    srill     ill    lie    eliitlelial    eelU    1 1 1'    til"    intestine    al'tiT    t'attv     fouil 

lias   I II    t.'leii.    wliile   tlie\    ale   aliseiit    .|iiriii'_'   slar\a1iuii.      'I'lie  e   i)af 

tieles  seeiiieil  to  liaxe  iia-.-e(l  iliiiM'ily  ri-niii  tlie  intestinal  eaiial  inln  tin' 
epitlielial  eelU  lieeaii  e.  wliiii  llic  I'.'t  was  staiiii'il  \\itli  eliarai'ter'slie  fat 
stains  liel'ine  ri-.iliin^  it  tn  tie  .■■nillial.  tlie  irlulmles  in  tile  e|i'llielial  eells 
wei'e    IniiiKJ    In   I"'  siiiiilavl\    stailjeil.      'i'lie  sii|'|iorte|-s  of  tiiis   liieelianist  ie 

\ie\\   nf  I'al  ali-iii'iiticin  iiiainlaineij  iliat  tlie  apjiearan f  the  stained  fat 

irldliiileN  in   ll jiltlielial  eeil-,  eiiiilil   in't   lie  exjilaiiieil   ill  any  ntlier  \va\ 

tliaii  li.v  siiiijinsinir  tliat  tlie  fat  ijlnlniles  lunl  uanijereil  iinhripkeii  iiitn 
tlie  epitlielial  eills.  Siieli  a  inneliividii  is.  li.iweviM'.  iiiiwarrimtecl,  fur  the 
stains  that  aie  xiliiMe  in  I'.it  aii'  aKd  ^nliiMe  in  soap,  mi  that  when  the 
t'al  splits  lip.  the  slain  w  i'l  leiia'n  attached  In  the  snap  and  lie  carried 
alnii'-'  V'.  ith   it   intii  the  intestinal  epilherniiii. 

.\-lis,,|iil.'  |M.  nf  thai  the  elieniieal  Ihcvy  is  the  ei^rieet  one  has  heell 
supplied  hy  a  lai"e  nniiilHT  of  r\  peiiinenls.  The  follow  inu'  may  he 
citeil:  I  Whc'i  Ih.'  iMiiph  llow  iio.;  I'l  oiii  the  thoraci<-  duet  is  examine  I 
after  fi'i'ilii'i;  with  latt;.  acid-  ii  stead  i  f  neutral  I'at.  it  is  found  to  coiitiiiii 

oiiK    iieiilial   fat.  iiidiealiii';  liiat   .'i   synthesis  must   ha\ •ciirreil  hetweeii 

ulvceriiie  and  fallx  acid  iliiiiiiu'  the  ah -oi  pt  imi.  The  '.dxeer'nc  for  this 
synthesis  is  furnisheil  fi  oia  soirees  which  will  he  d<  sciilied  liiter.  ('1'^ 
When  an  emulsion  made  paill>  of  neiitr.-il  fats  ami  pai!l\  "f  some  li\- 
ilrocailioii,  sneh   .is  a'helcMc.  is   fnl   and  the  fece,  are  cNamini'd   for  tliese 

siihstaiiei^s,   it   has  1 n    found   that   all   the    j'at    hut    none  of  the  livdi-iicar 

lioii  is  ahsorhe.l;  the  f.'c.s  contain  ali  of  the  all  oleiie  liiit  none  of  the  fat. 
This  e\pei  iiiii'iit  supplies  \ei\  stroll'.'  evidence  a'iaiiist  tln>  mcidninisti- 
thenrv,    for    microscopic    examination    of    the    ahove    d-crihecl    eiiiulsinii 

shows  the  partii-les  oi'  mulial    fat    and   li>  diocarl to  he  ot'  exactly   the 

same  si/c.      ■:!     I'.y  examinin<_'  the   piopciiics  of  the    fatty   suK-ta 's  in 

the  ilicuacie  Iviiipl Ilccteil  diirini;   llie  ahsoiptioii  of  siiidi   an  emulsion 

as  that  ih'scrili-'l  aliove.  iiolhini;  hut  neutral  I'at  li.-is  h  I'n  found  pi'esciit. 
i4  ."similar  results  are  ohtained  when  wool  fat,  which  is  an  esli'r  ol 
idiolcsliTol  and   fatt\    acid,  is  WhI. 

\Vc  ma\    ciiiicliidc  th;it    l\ill  i  siiliylnidi  a  iihiili  iiri    iiisnlii}ili    in   nnlir  or 

run     Hill     III     lliliuliil    Inl    illill  si  mil     mill    silh\lil  llri  s    Isnilji)     Hull    illi     ,s'i//)//l/. 

///  iiiili  r,  iiri    mil  iih^iiilii  ll,  liiiin  n  r  hi,  i   Inl  Ihi  ii  iiniii  III   III  iillii  r  i>iitl  irulnts. 


FAT    Ml  TMinl.IsM 


no^ 


TliiM-li.ii.i.  .il  Ili.M.iy  of  Till  iil>snr|.iii.n  rurtli.T  ,  Npl^iin^  wli.v  lli-iv  slioiiM 

1,..  Mir).   I;..-.'  .i.iMMlitir^  of  suapy  M.l.vtM. ^   n,   \\r   int.>tini>l   .Mmtciils. 

..|M,1  ills.,  uliy  tlM'  L'li.lmh's  ,,(•  C;,!  ). ITS,. Ill  in  tlif  .■i.itliflial  .-.-lis  ,.f  ill.' 
inti'.tit..-  ar.'  sn  viy  iiiiirl,  siiiall.T  lliaii  llm...  u  liirl,  lir  .  n  tlw  smfii.-,.  ..f 
till'  t'|iitlii'liiiiii. 

It  im'm.'IiI  1m.  ol..i-,-!,.,l  t,.  111.'  ,M.n.-lii-.iM,,s  .juM  statr.l  tluil.  iiltl.uudi  lui.l.'- 

,,.,.t„Mr.  tl.-r.'  is  ivally  s,„ .s,.„tial  pli\si,-al  .lin'.T.' l..'txv.'-n  .'innl- 

Mli.Ml  fat  aii.l  riMulsilicl  liv,lrn,-ail».i.      In  onlrr  riitinly  to  |«r"V.'  tin'  .'Msf 

,■„,.  tl..'  rliriniral    III y.   it    is   i s.„ry    to    \W,\   a    h.^lltnil    filt    ,.uss..s.i..<.' 

s„ •li.M-a.-l.'iistir  tliat  ,1.m„.ih|s  on  tl,.  iiianncr  of  iniion  ..xistin-  l.-twrcii 

fatty  a.-i,l  aii'l   uK -rriiir.  .-in-l   tlirn   to  >,■,•   uh.lli.T  it   aiM»"ii-s  in   an   mi- 
..lian-iMl  ron-lilioi'i  in  tlir  llior:i.-r  'I'l.-t       If  it  ilo.s  s,,.  tli.'  fat   must  liavc 

1 1,  al.sorl.-'.l  tlirou-li  tin'  int.'stinal  .i.illH.liniii  in  an  nnlnolv.ni.  nnsapon- 

i(i,..l  ron.lilion.  for  it   is  nnliK.ly  llial.  in  tl,.'  r.syntlnsis  u  lil.'!.  ...■.•iiis  in 

tin'  int.'stinal  .•pitlicliuni.  tli.'   fatl\  a.-i.l  nml-'.-nlrs  uouM  i uiiImm-  with 

till'  }rl><''M-iii('  inol'-.Milcs  in  .■Na.-tl\    tl,.'  saiiii'  iinmi  .■!■  a-,  lirfoi.-. 

Tli."v  aiv.  liou.'V.T.  hnt  \.iv  f-"  .iiialltir,  of  n.'iitial  fats,  at.art  fp.ni 
tli.si'  i.niir  falt\  a. -ills  wlii.li  .•oni|«os..  tli.ni,  l.y  vvliii'li  lli'y  '-an  Im'  .-liar- 
a..t,.fi/.'.1.  Til.'  nn>-1  liU.'ly  .mi.'  is  tliat  of  ..pti.'al  a.-tivity.  No,,,,  of  tl,.' 
o,..lii,ai-y  fats  is  .,|,li..ally  a,ti\.'.  alllion-jl,  fn-iii  .■l,..ni'.'al  ...,iisi.l..i-atio„s 
it  is  .|, lit-  .•o„....ival.!..  tiat  s,  ni..  s|i.,nl,l  !..•  so.  In  .,nl..,-  t..  .,l,tain  s,i..|,  ii 
fat  |!l. ..„■'■  i-,.,,.!,!..!.'.!  iimii-.rons  .■N|...ii„,.'nts  with  tlio  .'st.Ts  .,f  s^'ari.' 
f)..i.l  *  li,  a  s,.,i,'s  .,f  ..\i',.,M,i.',its  111.,,,,,  fcl  is.,)„a,iir'(I  .lilamat.'.  ii  syii- 
tli.ti.-  fat  .,f  .l,'M,-.,n.tat..,y  ii.iw.T  ami  as  ,...a.lily  alsorln  .1  as  natural  fats. 
.,,,.1  l,y  .■\a,..inati..i,  of  tli..  inntial  fat  nivs,  ,il  in  tl,.'  .-livl..  ll>"in'.'  fn„„ 
H„.  tl,o,.a,.i,.  ,1'iH.  h,nn.l  no  ..\  i,!'-,,....  «(  \h<-  .l..\t  ro,',,talo,-N   tat.    Tliis  ,-..siill 

ronlirnis    |„...vi,,iis    w.,i-k    l.y    Frank,    wli..    fo 1    Unit    ll'.'    .tliyl    .'sL-is    of 

fatty  a.'i.ls  a,-.'  n.,t  al- .h-Im'.I  iin.-liati'j.'.l.  Tl,.'  I'.'siilts  .,f  l„>tli  «.nl<.'is 
onip'liasi/,.  til.'  i.,-.,l,al,:iit>  tlial  ,',-i.lil.\  -a  p,,i:itiaM.-  fallN  a.-i.|  .st.Ts  .1,. 
not  .'si-ai...  saiioiiili.'ati.,,,  \in.l.-f  tli.'  fav.naMc  .•oii.|iti..iis  of  lli,'  n..iii,al 
Int.'slin.'  In  otlnr  «..i-.ls.  In,, I  lli,'  fat.  K.'.'n  al.s..,-!..'.!  iin.-liaii'j.'.l.  a- 
M,,iil,|    I..'    ,-.',|ui,i-,l    l,,\    tl','    ni.'.-lianisti,'    tli,.,r>    of    f.it    al,s,n-i.li,.,i,    tli.'y 

v.Mll.l   lia\,'  app.'ai.'.l    in    tl lix  1-'    in   op' ''•••i'l>    •^■■^'^ n.litions. 

Tlii'si'  iin.M  impoil.ml  .■oi,.-lnsi.,ns  l.a.l  ns  to  in.piiii'  as  to  //'.'  r.  i;.s'"» 
f„r  tin  rinnnj.  ill  hit  ,hnn„l  >ts  „hs,.f,,l „,,,.  It  ran  not  Im'  f,.,-  til,'  pui'pi.s.' 
,,f  pr.'v.'iilin-  111.'  al.s.n-plion  ..f  nn'l.'sicaM.'  fatty  -.iil.stan.'i's.  sii.-li  as  lli.' 

p,.t,'ol..,in,    li\-l,.M-,-,:l",ns   .„■    til,-    u,,o!    fats.    1 ans,.   s,,,-!,    s,il,stan.-s    a,.' 

s,,  ,ai','lv    |.r,'s,.n1    i,i   ,,iii-  f I       H    i-  "i..s1    p,',,l,al,l..   tliat    tl,..  ln'..aU.I.n\  n 


•|..l,„,r   |.!.--i..,| 
;    |,.n,.     .,,..1    I.,    I,. 


,, 


l„jl|    ,11.  ll 
-!,.«     I 


..    ,„.,1U     .,..,>.      ,„..,.,'  •..■'    •!'-'•■,,  ."       '  ■■'     I 

,      111.     ,-   ,l,t.  ."II,     .,>   111.    •■,  !,,..,i     '.,'^       ll-    ••.'-' "    "■'- 

;;;-;;;•■:;•  ;:,,,;::,:;^.::,^;r,.;t':.;.^.•::v^/:':.:::i^^.:';.■-:^^^ 

,l,.-,l    aJ.,,,! ,,,..       ....    -'    ...    1... y,s     ,„.„l.,,i      ■.^.,.^,.,.„.„^     ,,,..,.,.  h.  h.    ...'. 

Mm-    t:it    •..■„>., I     .,,,1,11.       l!.,ril,ir,.    Ui     !.-l,i;    11,1"    ..     ^.      '1  ,"'■"" 


ll„«     1,, 


,it,     -:,J„,t,,.r.  .it,',,t 


sv  ' 


i  '. 


J  1 


I 


Mi 


p  I 


^i 


ii 


L^m,t  If 


I'. 


I  f 


i  It. 


i 


ii  ^-i  i-si 


«  •■:.■   ! 


ij 

i 


f  f 


604 


Ml  ■|\r.nM>M 


illKl    tvv\lillir-is    ..f    llrUll:ll     Tilt    (■(•.•111-     I'nl'    ttir    -.•lliir    lv;l-,..n     Hi, it     MMllLir 

l„-n,-.'«.s,^   ur  (ImiiiL-    tlir   ;il-.iipti..n    .-iii.!    n^-'iiiihil  i.'ii    "f    pint. 'in,      '' 

Mill    |„.    ,,.ln.lill.,T,vl    Ill.lt    |.l..t.'i||    is    .■lllitvl)     dlMlllc-^lMlril    ill    til.'    illt.'stilH 

i,,t,,    it.   -,,  ,.a!!r,l    liiiil.liirj    -I .V      TiM-r    .-.iv    ;,l.-.nilK,l    s,|.;ir;itrly    int.. 

til..    1,1 1,   ulii.h   iMi-ri.'s   tl,.in    t..   tl,..    tisMirs.    ill    wli'.-li    tl.i'V   I r.' 

sMitli.-i/.'.l    1..    l-iiii   111'-   l""l>     j.intiiM.      An.l    v..    11    .iiMM'.ii-   1"   !"•   ill    ''"■ 

.-.•iN..  ..f  tilts.      Til,,   pr vs.   in   ..lli.T  w.if.N.   |...iiirns  .,r  tin-   i.'iilliurj.-li t 

,,f  f;itt>    .-i.-i.l   lll..l.-.Mllr,.  ;is  ;i   ivsiill   ,.\    wlih-li   the   li.'«l\    I.TliM''!    I'.il    i-  1 ■'• 

;i,!„pt.,!,|,.    f,,r   u-r   ill   li ruji.isiii.      II    1 1'-   I"   ''■   11.. 'iv    liK''   ll ''■II- 

jirlrfisti,.  t'lit  ..t'  til,,  iiliilii.-il.  TImK'  IIKIV  lie  ;!  1 1 .  •!  ll  .M'  IViisnli  tnf  Hir  pi'"- 
,.ss  It  \'  ill  !.'■  f. . 111. .Iiil.. .!...!  Il..-,t  l.-.-itliii.s.  \\lii.-li  (-..lislitllt..  til..  III. .si 
il,,p.,i-t.-nil  ..f  111'-  f.itly  siil,st;iii.-..s  .,f  ll, II   its,.lt-.  ,-1...  ...iN'.l  .jlx.-.  li.!'--- 

tll.-lt     is    !,,    s;|\,    ,.11-.'    .•..|llp..||ll'!s    .•..|i!,-.illil!'J     11     \alilt>      '.r     lillt>      ilCl'i-s.        Til.- 

iv.iiTaii'j.-iii.-nt    ..f    tl;.'    iiii.!.-.-iil<-   "I'    ii'-nti-al    fat    v,  lii.-li    ....-iii-s    .liiiiii-   al. 

s..ipti..n   iiia\    1"-  ll;.'  tir-l   -l-'P  in   ll"'   1 1  :iii-l"i  ni.i' i l'   ''•"    i'""       '-illiin 

111  .inl.i-  (.1  llii-(.\\    lui-tlH-i-  li'jlit   "11   111'-  .|ii.--t!..ii.  Ill ■   :  ..>  1"      ■'fill'    ' 

ji  iiiiMil.'-i-  ..f  int.!.  stiirj  r\p..|iiii.  iits  in  \\lii..|i  th..  fli.nii.Ml  |ir"|H.|-t  I's 
,if  t'.its  l.,.f..r,.  ai'l  .-ill.-i-  a!.s.,|-pli'.ii  w.-ri-  .-..iiipa  i  .-'I.  Tli.-  i-iit-i-i.-i  «lii.-li 
ll,.  t....i.  wi-i-.-  iii.'tiii.j  p. .int.  i'Mliin-  \:ililr.  an.l  Iih.im  111..I.  .ii!.-.  r  w.Mylil: 
til.,  iii.llinv  p'.int  r.-pr.  -.  nliii'J  tii.-  s,,li.|it.\  ot'  tli.-  I. it.  an.l  tli.-  i.Mliii.- 
\alii'  .  its  ,1.  .^1,  ..  ..I  ii!is.ini;.iti..n  lli,-it  is.  ili.-  iiiii.il..-i-  ..f  .l.'iil.lc  Imlvs  m 
til,.    t'.,tt\   ;;.-i.l    i-liain        It     \>  as    t'..iin.|    that    ijiiiin.^    al.s.,i-pt  i.m    \.m\     run 

si,|,.i;il.i,.  .liaii'ji-s  Ill-   111   tills..   n\.,  .-liaLM-t'  listi.s;    |-,,|.   i.\aiiipl'..   "Inn 

fat  uiili  l.i-li  iii..|tii|._'  pi.int  .111.1  l.iu  i...lin.  v.-ilm-  \'.as  l'...l,  lli.-  lat  in  lli'- 
tli..iM.-i..  IxMipli  Has  .,|  .lisiiii,-ll\  l..\\..i-  iii..|tiii.a  p..iiit  ami  lii'^li.-f  in'liii' 
\alii.-  Wl.n  tat  \'.  itli  a  !..\\  iiii-ililiu  p'.int  an.l  a  in'jli  i."liii.-  \alili'  was 
|-,..|.  til..  >.-,ii-s..  i-Imi  •^..  ...•.■lii-f..!.  f.T  till-  lii.-!liirj  p..iiit  "I'  till-  lli"l-a.-i.- 
l>liipli    ''.il    ■    .is   lii-li'i    ai.'i    111.     i...|iii.-    \.i!ii''    l..U''i-.      TIl-si.    I. -suits   c.iil.l 

1 \plaiii...|   .IS  .In.,   ill   tiM.   liist    ,.,.|s..   1,,  til..  ..i.l.iilii.i!   Ill'  ..I'-:.'  a.'i.i  t.i  'I,,. 

fat    .Iniii"-'    lis  sx  iitl  .sis   111    till.    iiil..siiiial    i-pil  |...|nilii.   an.l    in    lli.'   s,....,n.| 

IMS.,   t..  till.   .i.|.|ili..!i   .it'  s ..iiiiiati  .1    t'ativ    ,-ifi'l- 

Wh.M    a    111 slsnii.j    ii,,iiiil\     ..I'    ■j|\....i-|.|.-    .111.1    satiH-al.-.l     TaUx     a.-i.l. 

I, lit   \x  ill,  ,,   I..U    iii'-ilii,.-'  p.. int.  »\as  r.  .1,  ll  .    a.l.li'i.,1,  >•(  "l-i'-  .'I'-i'l   ^^as  still 

!,,iin.|   t .111.   .IS  ,ii..L"-'l    M-..111    tli.-    M.ilili.'   \.ilii.-.      'I'liis    ill. I,.., It. -s   that    lli'- 

,.|ia,|.ji  !-.  11. .1  iiii:.';.  in  ..nl.-i  tiial  llf  iiirllin'^  p.iint  ..T  tin-  al.s.iil....l  lat 
„i;,.,    |„.   I, ,.,..... I.   I. Ill    .il-..   I'.. I-   s,.|ii..   .■liiiiii.-.-.l    i-..,-is..n.      In   a    f.'iiill,   s..ii..s 

..I'  ..\pi-iiiii.  Ills.   .,    |..\.  .iii.j    ..I'   i..'rini-    \a!ii ..Ill  11. 1    all.  I-    t'.-.-  liinj-    "itl. 

,.,,.|!i.,.,  ..  .,i!.  v.iii.!  .•..,,:aii  s  a  ln-'li  pn  .-..iil  a--'.-  "t  'jl.v  .•'  li'L-s  ..T  ITrjlilx 
lllls.lt  1,1  ..,1.  .1    t.:lt>    a.-i.l. 

|.;.,  i,|,   n||\.    111,.,,.    //   .      ,„l,.t,,,,     l„,..,  ~s.   .    U.,     i,ni,,,-..l     „n„l,',lnl,l    fl,,     r,,„l 

l„i<iluni  ,,i  f,!>  (hinii'i  il.<  .ih~.,ifli"".  an. I  tins  iii...lili..iit  i.'ii  is  appiii.  nth 
n|'    siii-li    a    liiiini.-    lliat    it    rails. s    a    i-li.iii'jr    tuxai'l    tin-    pi-...iil.-l  i"ii    "I'    a 


\MU 


K\T    Mi  I'M.'H.I-M 


(!!>:» 


imit'oiiii  I 

IlilliL''" 


li\  li'    fat,    |ilTNiiiii;ililv    chiiin 


•  ti'i'istx     "!    Hii'    ;iliiliiMl 


Mfi'   i>niliiilil\    'jMiilir   than   '•«.! 


t|i,.  iilisiiilM'.l    I'.-it    ]ifc-iiit    in 
ill,-    i.lcic    ariil    III'    nf    till'    ^a 

•Ni'iit  \iiiUn'i\\  II. 


ll.l    l,r    |,r..,|ur.'i!    1>>     ;Hlliii\tm-c    »\ 
•livl,'.    liUl    till-    -..ui-i-i'   "f 


tlir   imniial    tastiiii 
tnrati'il  a^-nl    iri|nii-r'l    fuf 


this   >\  iitli.-is    Is   at 


lUi 


il  ii  l 


!       il   '. 


u 


iiit; 


iil.i  i^ 


jl 


11^ 


1l  .i       I 


CIIAI'TKI,'   I.XXVIII 

lAT   MKTAI'.til.ISM    iCuiuMi 

THE  FAT  OF  BLOOD 

Methods  of  Determination.  Ninnijillv  ilir  M'hm!  ci'iitjiiiis  .mlv  a  sni.ill 
lirii-i'iit,ii;i'  1  l'  t;il.  I'lil  Ml'lir  ,i  I'.ittv  \i\-n]  it  iii;i>-  riiiit;iiii  ^n  l;irir<'  mii 
jiiiiiMint  tliMt  till'  r.-it  .-M-lii.-ilIv  ri-rs  to  llir  Niiri'iiiM'  di'  tln^  IiIu.mI  HIm'  a  i-i-isim 
l!y  iiii'iiiis  lit'  tilt'  iiltiiiiiiirrD-M'iiiii'.  cxaiiiiiiMtinn  o|'  tlic  Mimil  in  tlic  ilark 
lii'lil  at'tci-  a  I'at  lidi  iiiral  ft'\ca!>  tin'  pri'sriii'i'  ni'  irlaiiciiiii'  |>afl  idr--. 
till'  M>.-all.Ml  '■U\\  iliist."  'I'lii-.,.  |Miti<-li-s  all'  tiiii^l  aliuinlaiit  almiit  •^i'^ 
limns  al'tiT  tin'  im-al  lia>  liri  ii  taken,  ami  tiny  u'l-ailiially  ilisaiiiicai'  1'" 
till'  twi'll'tli  liiiiir.  'I'Imv  ilii  iii't  appiar  al'trr  a  iiii'al  wlii'ii  tin-  tlmfarii- 
ilili't    i^   liyalrtl.      'I'1h-.\    ili--a|iprar    w  bni    nW'.Tii    is    IhiIpIiIi'i!    tlirmi'jli    tlir 

lilui..]^ 

|-'al   iliist    lias  alMi  ln-'ii    ruuinl   aliiimlaiitlv    |iii'sriil    in   tlic   IiIiumI   nf  riii 

lit'.vii  Liiliiii'a   |'ii;>  at    liili  tiiiii',  liiil   imt   in  tin'  iimllii'i's  M 1.     'I'liis  wmil'! 

iinlicalc  lliat    tin'  placi'iila   iniisi    lia\i'  llii'   pnw  I'l-  ni'  takiiit:  tin-  i-mistitii 

cKt-i  iif  fat    III  111   till'  iinitlii'i-'s  III I   ami   iMiililiiiir  tlii'iii    inlu   I'at.    w  liirli 

tlii'ii   passes  intii  till'  lili  11.1  nt'  llh'  iVliis      Till'  plai'i'iila   iiinlir  tlii'^i'  nimli 
tiiiiis  arts  liki'  till'  iiiaiiiiiiai>    ulaml.      In   this  ci.mmi  rt  inn   it    is  nl'  iiiti  ii'sl 
tliat  tlirii'  is  iilsii  mill-. I  I'at  pii  siMit  in  tln'  lilni.il  ul'  pii".i-iiaiit  wmiii'ii.     Tin 
I'at    I'liiiti'iil    111'  till'    plari'iila    is,    1iii\m-\ n-,    uieatn-   in    tin'   rai'ly    slaiii's    nf 
l)ri"_'iiam-\    tli.'in   latiT. 

.Mttinlltrll     tlii'si'     I'ai-ts    lia\r     1 II     klmwii     I'm'    snnii'    lilili'.     it     lias     lii'i'M 

iiiipiissililr.   rilliii-  mi    ai-i-minf    nl'   tin'    lai-'M-   i|ii;int  it  irs   nl'    lilnml    rripiiri-'i 

t'nr    a    i-lii'liiii-al    i\aiiiiiiatinn    m-    l ailsr    nl'    tin'    ilil'linilty    in    rst  iiiiatili'J 

tin-  anmniit   nl'  I'at   t'lniii  tlir  i|.'!isi)\   nl'  the  •"i'at  iliisl,"  in  riillii\\    w  itli  an\ 
iri-cal   ili'i,'!' r  aiTiiiai'>    tl \,ii't    i-ln'inii-al   i'lian',;i-s   tliat    taki'   plan'   in 

till'      I'at      III'     IIm'     Mnnil  U'll'intl>.     IihAi'ViT.      I'.lnnl-     |i,|s     sUl'I'l '1  i  I  I'l  1      ill      I'klli 

nralili'-'  nii'lhnils  li\    \v  liirli   tin'   I'at   ti'llt   nl'  ill.'   lilnnil   call   In'  i  Irl  iTIniln'il 

with   satis|':n'lni\'   aii'iir.irv    in    siji;il|    ipiaiit  it  ii's   n|'   lilnini.   sn    that    a    i-mi 
liniiiiiis  si'i'irs   111'  nlisi'i\  at  imis   i-an    In'    niailc   n\i:'   .i    i-misnlrialili'    pi-iiml 

■I'liv   f;il   i-  ,\|i:,.i,.!   I'm. I, I   til.    I!.-. I   l,v   : lr,.||..|  .  i|i.  i    tnixt.jri'  with   rii...|i  rate  In  ;il. 

An  :ili'i  i"i   |".i '  I  ■!,  ■■(  ll.r  till  III!.    (-  .  \:<\  •.lat.'.l   ill  Hi.'  I".  -.11 r  -.■.liuMi  1  lliU;il.' .   «lii.  li 

^:i| Ih  -    lli.     l,a.      'I'll.-    r.-iiii..    ...n^i-tnu    "I'    -'•■*]•.    i-    w.H    »;i'lii.|    :ih.|    tli.n    li.-.-i(.l 

Willi   !n.|r-.  M. •11.    :i.;.|   -..  ;,-  I..   I  1.  .  ipil;,l.    tlir    l^.ll\    :i.|.|        'I'li.    ,1,  ii~i|  i    ..T  t  li.-    |it..  ipil:.'. 


!    ■   ' 


■■•AT    M I  r Mini  ISM 


(197 


III,,,     , |.i.-,.|     is    r..,i,|,;,,r.l     ill     :n.    i.|ti,  1,1     i,|.|.;,,lilu~.    r;,llr.l     li     )„  pl,rl..|,Ml,T,     uitll     :i 

•Mii.liml  M.lnti..n   ..(  Ih..  inilli- imiiw  ,.(  ■■].„■  ii.  lI   in;,i.'l    iii   liir  -;i »:i.v.     Tl.r    f:ilt.v 

;iii,ls  ill  liiiiiiNii   I'l I  iiM'  iii;cihl\   'Ii  ic    ;iii.|   |.iUniili.-. 

Th.'  Irnil.iM   ;iii.l   .li..).-!.  n,|   ,    .,»    :,l- -liiHrit.M   in   lU.-  <,w  1  1.h,.1  ..xtniH.      For 

I,,.  11,1,1  tlir  iil."x,   lAli;,.!  ..I   1  I.,    i.  .111.1   ill.'  1.1I1..',    1  ..r  III.-  :il."l."l  liMil  I'tliiT,  is  l^l;;l■^.t.•.l 

1,1  li.iiiiiiu'  vMl'i  ."iM.iiiiMi.a   UN",  I  II  >^'>,.     'I'lii~  'I n|"-"»  ll"'  l.'.'itliiii.  liLrnitiiiL' 

ll„.  |,|i..,|.li.iiii-.  :,  >,.liili r  11.    t.-i!liii-  :i-li  l.iin-  nn'l.-i.'.!   fiiiiitlv  ;ilk;iliiu'  I"  pli.'!!"!- 

(l.tliuliiii   iiii'l  'I..  11  -1.>»I>    ^I'l'i"'   I"  ■■'   ^il^'i    "I":'!-  -'l"!!..'!.     Til.'  .I.n-ity  ..(  111.'   pu- 
,,|.it:,t..  thus  i.r...|.i..'.l   I,  ...II  |-;.i..l   .'i   H"'   ii.  pli.  l-ni.i.i    "iHi   Hiii'   "I'  "    I'l' -Mil:"''   I'r'" 

,1„,,.,|    ,„    tl„-   MUn.'   1.1 lilt    ■■!    -iImi     I  'I'M.-    l.v     ...Mill.;    !..    ii    tl    -C.ii.l-.l.l    |.l..,-|ii..lir    ll.'i  I 

>..|iiti   II. 

|.-,,,    ,;,,.;.,«',,./    .,.,    ;,li.|ii..l     |... 111. .11    ..I     III.     iil.'H.'    .Mi:..-!     i-    -:.!•■  i"li.-l    "itli    >...lilllil 

,.,|n|:it .!  'l"-i.  si.liiii.l.il  "I'll  .liliiiii-Mi,  ;  III.    .1.1 !•  nil  -Miii.t   .-  t,    \,  .1  uitl I  i.- 

,,„|u,lii.l  iiii'l    HX>.   'ill'. I  111.    lil'ii-li  .1.1.1.    I-   r.illv    .|.v..l..|..-.l      l...-1-iiiiiiiin   r..;in- 

li..ll_l.     111..     Ilil..|;-lli      ..r     VVI.I.   Il      I-     'lii-i iplil'.  'I      •' 

,,,■   li-i.    !■.;,.....  I.!    <' :■    -I..1.I1.1    '     L.  l.-l.-ml    -.'I'lli 


.|..iinii  1.  1    uitli   tlt;it   ..l.lnin.'.l   Ly 


Variations  in  Biood  Fat.     In  il..'  'l-^  il"'  i"  i.-'-i,i.i.^..  ..i    i;it   in   tin' 

1,1, M„l    is    |-ilii;itk,-i!.l\    .•uiist.-ii  1    nil. In     ii.iiiii;il    > Iiti..ns.      .\!l,r   «    f'ltl'l 

„,,„/  i|„.  ii„.i..;,s,-  iM  Tilt  l-.'.^i.  s  iii  :il.,,nl  tin  li.iur.  :iih1  iv:i.-Ii.'-  iN  tiiaxi- 
„nnii   111  .-iliont    six.     Tli.'   iii-i.;.,..   is   n.n    !..iin.l   in    aiiiiti  lU   •      ulii.-li    tin' 

i|„.r;i,.|,-    .liiH     liiis    I I,     li.^,-il.'.l        Ailli.iuuli    till-    ii-nl'     «"i=i'l    -■"II    '" 

,.,,„traili.-t   III..   M.'»    lirl.l    li.^    •'■'I'-'  lii.i'    I'.-"'    "1'  "■'■   '■■''    '■'''"■''   '•'"'    ""^   '"' 

;i ,,i1,.,l    i-,,r    ni    111..    ilM.r.-i.ic  .!m.M    I.Miipli    i-   iil'-'iil"''i    1'.^     ".i.^     "'    ''"■ 

,,,,,.,,1  vrin,  il   .I."'-  Mil'   I  >    i'-'ll   .lis|,i-,,,..  th.'  liM.i.lli.'vi-,   I'll    !'    liii-  !"■'•" 

|',„,H,|  ll,;it   tl,.-   Kil   I.  lit    .11    tli.     i...rt;ll   1,1 1   is  :ilu,i.\>  lii'jli.M    tliail  tlliit 

i.r   tlir   .jil'-'llliir 

V.Ty   iiit.Trsliii.j   r.-siills  l,.,v,.  I I!   ,,l.l;iin.'i|    fiillnu  in-  tin'   n,fr,fr,n,„i^ 

n,;,,i;„„   i.r  .Miiilsi,.!,-  ,,f  .nl.  ritlin-   ih.'  s,,  .■.■ill.'.l   i-Ms.in   ,iiiiils-,,n   ,.,!■  .•,,!- 

i,,i,i,.,i  His|.,.iisi,,iis,    rp  I..  II  .Ins,-  ..r  (•  ♦  'ji-.-iiii  |H'i-  Liii.-i-;nii  ..r  iimiv 

xv.-i.^lit      Mhi,-li    li,\    .•al.-iil;iti..ii    \v,,iil,l    snflii-i-    tl,    r.-iisi'   tin-    lat    ti-nt    "I' 

t!,,,  1,1 1  |,\     |iiu  |n  |.  ,-,.|it      lli,.i-..   was  11,1  in.-i-.'as..   in    I'at   .•.,ii1i.|it.      In   <ii- 

,1..|-  In  .-vplain  tins  ,|is,,,,i„.ai-.in.T  »\  lat.  it  nu-.-lit  In-  iiiiauiiwil  th.it  tln- 
iuj,.,-!,.,!  r,ii  parli.-lis  i,,riii.-,|  ,-iiil...li  in  i  In-  -iiiallrr  .-apillari.'s.  A.jain-I 
sui-h  .-1   \i.-u.   linu.-M'i-.   is  111.-   la.t   that    til.     |,arlii-l.--  .'f  lat    in  tins.-  .-itiul- 

si,,iis  ar,.  nij,'  liair  : n-  s..\..nlli   tin-  s,/.-   ,  \    a    i.-.|   |.us,-!...      AlllmMiili 

this   aii-niiinil    is    |,,.il.t    ..T    -,,„„■    u,-;.J,t.    it    s|,,,ul,|    I..-    i  ,-iii.-iiilH-n-.l 

tliat  III,.  |.iivsi,-;i|  ,.,,ii,|itniii  111  this,.  Ill,,-  l.-it  .J., hill. -s  is  II. , I  thr  sain.'  .'is 
that  nl  111.-  II'. 1  hhinii  <-.ir|'iiM-li-.  Tln-ii  sinla.-.'  .-nii.liti.iii  may  lii.  su.-h 
that  th..y  i-.a.lil.v  a^'jliit  iiiat.-  s,,  a-  1..  Liini  Minill  iiiass,.s,  whi.'li  ni;i> 
sti.-k  at  III,.  I.raii.-hiinj  <•(  th.-  si,i,-ill,.|-  art.iinl.-s  an. I  ,-a|.illari.-s.  r.l.i.n- 
lniMs,.|rsu._'..'.-sts  ihal  till-  in.j.'..t...l  fa!  iiia\  h.-  s|.,|-...l.  (...ssihlv  in  th.-  liwr, 
sill,-,,  ih..  lal  ill  liiis  ,,i-.^aii.  .-IS  \M-  shall  s,-.-  lat.T.  in.-|-cas,.s  ninh-i-  siniil.ir 
.  iin.liliniis      \Vh,-n  lui.-.-  th.-  ahnx--  .|iiaiilil\    was  I'.-.l  in  the  r..rtii  >•(  .-'i'-i- 


il^i: 


V. 


f  :,' 


i 


i 


1 1  i 


ll',. 


cm 


Ml  rvr.Di.isM 


II 


.5' , : 


I  ' 


I    ' 


ynlk   I'.il.  M>nif  ..r  it  |..M>i>lnl   in   llu'  I'l 1   I'-f  M'vnal   Im.uis.     This  in 

riT.is.-  l.lJiy  liavr  Iwrii  ..uin-  t..  tin'   ll lillSi  d"  tli.'  t.'ln|M.rill  V  NtnlvlicMls.' 

xvilli    r.it.  ..r.  iii.il..   pnil..ilil>.  tn  ii    ivtiinlinu;  inllihiir..  tl.;it    Ir.-itliin   iiuiy 

1,,.,\. f.il   iis-.iiiiilMli..ii.   f.-i-  Ircitliiii   itM'ir  iMTsiMs  in  till-  I.I'mmI   f..r  a 

Idiij:  lime  altiT  iiitiavtii.iiis  in.jcclii.ii. 

Dmiii..'  fnslimi.  iiH  in,-ivas,.  in  l.loo.l   fat   uas  ri.uii.l  iiiil.'ss  tli.'  aniniMl. 
hy  s|)r.i:il   r.r.liuir.  lia.l  Im.ii  sliilT..l   uilli  i-M'."-s  ,,|-  lat   pti")-  U,  111.-   I'a-.l 
in-  p,Ti,Ml.     Tlir   llpriiiia    ill    lliis  ••a^-'   iM.li.•a1.■^   lliat    Tat    is  h-iiiu'  tian- 
,,,,,,,.,1  ,■,,„„  ,„„.  phu-..  t,,  aMHtli.T  to  MTvr  as  luri  I'l.r  ill.'  starxin-  tissiirs 
.V„r.../;.s-  u.f   r-nii.l   lH  priMliic-  an   iiinras..   in   l.!c»..l   lat.      Ktln-r  pro 

,1 1   iliis  iiMivas,.  liniin!.'  tin-  nairc.sis.   «liriras  nmri'lnn.-  and   rlil-.i-u- 

f,,ni,  .ii,I  iiHl  ,1.,  s,.  niitii  alt.T  r.MM.\  .Ty.     Tlir  -■xphmation   -iivn   I'-.r  tlic 
,.tl,rr  .(Ti  ct  is  lliat  a  luiMuiv  ul'  Mn..,l  ami  rtinM-  lias  lii-lirr  solvent  p..wi' 

fnr  fat   tliaii  M'mhI  „K 'I'll..  I'splanalinii   f..f  llir  .•lil..r..|-.iriri  aii.l  nmr 

pliiiir  .•ir...-ts  is  tliat   a   .Mitaiii   ai.n.nnt   of  lMvak.l"Uii  nf  llu'  tissi ■lis, 

in  uliirli  lipilis  ;,,•.■  sH    fir.'.  siip.TV  .M  i.'s  lipnli  tlir  artioll  of  lln-s..  na  rcot  irs. 

Thr  M'mhI   lat  also  iMM-oiiirs  ..iioniionsl\    in.Tcas...l  in  alnnit   forty  lioriis 

„,■,,,,.  til,-  administration   of  plilo,lii/in.  and   on    tli.'  s,.,.oiid   <>v  tliiid  day 

;i|-trr    liir    adiiiinistiation    of    i>!iosplioi  „>.      Tlir    s| ial    si-niliran.-,-    .d' 

ll,,.-,.  f.irK  uc  sli.ill  .-onsidci-  lat.T  in  .•orincrt i.iii  willi  tin-  r.lal ioiisliip  "f 
til.'  Ii\.r  t.i  fat  iiii'tal".li^iii. 

jiy   .•..inpaiis-.n   of  ill.'   fally   a.M.l,   l.'.-it  liin.   iiinl   .•li..!.'st.T..I   I.'uls  -'f 

1,1. ,...1  ./»/•..../   I.il  „h.,„-,,i,u,:.  it   lias  l..,ii    f..iin.|   lliat   tlinv   is  ii  slrM.ly  ImiI 

v.T\     vaiial.l.'    in.T-as,.    in    latt>     a.-i.l.    a.-.-.^nipaiii-Nl    l.y    no    vaiiati.iM    in 

,  !  o|,.si..r..l.  lint   uitli  an   in.-nas,-  in   I.MMtliin.  ulii.di  \arirs   fioin    10  t.>  :!.. 

..  ,.,.nt,  1,111   .].,.■>  II. .t    niM  stii.-lly   parall.'l   uilli   lli-'  fatly-a.-i.l   in.Tc;is... 

..   pr.,l,;il.lr   Itial    this   iii.M-.a-.-   In    h'.-ithin    (•.•prrsi'iils   thai    part    id"   Ih" 

,,,il„..l    fMt    uhiidi    is   int. Ml. I.'. I    f.ir  iMiin.'.liat.'    nsr  in   thr   tissiirs   (  pa-.'.' 

Tii.H.      Til.'  inoiv  or  l.s^  iii.l.'p.ii.l.'iit    in.-r.'as.'  in   h'.'itliin   is  .,f  si'.'iiili.'ai 

in   ,'onn.'.'tioii    uilh    th"    fa.'t    that    in    many   patholo'-'i.-   .■on.jitions   of   s... 
,.,.,11, ,,1  |ip,.,ni;i  ll,,'  in.'i-.as.'.lo.'s  11..I  alViM'l  llii'  fats  .,f  tli.'  Mo-.d  hut  rathi'r 

tl,,'   lipoi.ls     i   .'.   l.'.'ithiii   an. I   .•li.,lisl.r..b.      S.parat.'  analys.'s  of  1,1 1 

pl,.,s|.!M  Mild   ^^llol.■  111 1  sli.iu    Ihr  i ii-'na-.'  of  l.'.'ltliiii   1. 1  Im'  niiii'li   nior.' 

i„;,ik,-,i    in    ill.'   .'.,rpiis.'l,>    Ihaii    in    Ih.'    plasma,    uh.'i.-as    the    fatty  and 
ini'r.-.is.'   i-,  .■.inline.!   I.i   lli.'   plasin.i. 

Tn  illn-liiil.'  soiii.'  ..f  th.'sf  p.iiiil>  111.'  follow  ill..'  tal'l.'  "ill  l"'  "1  vain.-. 
1,1  It  ,,  .|,,vMi  III.'  av.ia'.'.'  .listril'iiiioii  "\'  fatl>  a.'i.l.  l.'.'ilhin  ami  .'Imlt's. 
l,.,,,l  ni  iM'iiiial  indi\  i.liials  milI  in  .'as.'s  .,f  .li;il>i't.'s,  in  w  lii.di  .liscas.', 
,'is  Inis  l.c.ii   known    lor  l"ii'-'.  iIk'H'   is  mark..!   .lisliirliaii.-.'  ..f  fat    m.'lali- 

.lllslll. 


1  I 


VVV    Ml  TM'.III.ISM 
l'l....)l.   I.1I..1I.-   IN    NoUMAl.   AMI  IN    III  Mil  Til-    rn;SMNS 


699 


I  MI1.li  I    VmiKKATK  KKVt.IlK 

N.mMAt.     I    |„^,„.|.^s     I    i,iMirrts    |    ihmutk.s 
rmiTNT         ,.|i:,KNT     '    rmiiNT     l    niirUNT 


I  \\li..|.    lU I    I        O..V.I 


I'M    l.v    I'.l    .T 

'ui.Mir  111 I  i       i>:'.7 

■ri.liil    y.:V^     A.l.K-     I'I:i<im:i  I          •'•'■' 
V,„|.„>,l 

wiiui,.  Ill I  :       ii.:'.ii 

I.,  nil, in                               l'l..M.i:i  I           n.'-'l 

(•,„l..i..l.'^  !       ii.l:: 

-    \Vlr..I.     I'.l I  ".•.'•-' 

I'l  "I'^li  1"!                        .      |'hi«lii:i  l'.-^'' 


ii.s:; 

II.IMl 


i;!v.  li.l.  ~ 

I'mIi.I    l.i|".i.ls 


!  jC,,i|.>i^,l,,  li.U'i 

I    I'l;,-!!.:!  I  li.l' 

I    C.M.u-.l,-  H 


M.i;i 
I'.:;  I       I       II.  I.', 

ii.:;l' 
II -J  1 
I'.l:; 

ii.lJI 

ll.'JIi 

I'.L' I 

n,:s 
II. 1^ 


liM.-i,i,i:i 


(i.fis 


O.'.ll 
l.dti 

II  I'l.'i 
ii.T.'i 
(i.lS 

ii.;;:! 
I'.-** 
II. Ill 

1 1. I'll 
11. ".II 
I'. I'll 

II.  lis 
1 1.  •_':'. 

l.lii 


III 

I.MI 

1  III 

I.L's 

Olll 

II.  Ill 

0.  Ill 

on 

II..-.  1 

(i.'.M 

||.>>I 

ii,::s 

!.'.•>* 


I,    „i||    1 1,>,.|M,1    llml    IImI.'    i^    al.nllt    UT    jii'l'   .Till    nf    I, .till    iMtfy    suli- 

.,,,n.-.-  ill  M..iin.-,l  1.1 1      Tlir  l;in>    .■i.-i.l-  i|i;ilMiilii-  AuA  i.l.'ii-     Miiiiinnt  1.' 

;,|.,„„     in     |„r    ,-,.m!.    ,u,.1 .ill.^     ilwMll'Utril     I...Uv.'ll     l.hlMMM     iUi.l 

,.,,r|.UM.l,-,;   llir   l.'.-illMii,   Ml.niil   n,:;   |,,.|-  .thI.   l.i'in-   tuir,'   ,i^   ;iliiliiilMnt    m 

,.„,.|.,l>.-li'>    :l>    ill    PS.IMII.I.    I    tlH'    .•l|.'ll>t..rnl.    n.-J    |„.,-    r,-|l1.    illiiiUl    lllll.-'ll.V 

,|i.triliul-l.      In  iliH-.t.-.  Mil   nl    !l,.M.  Mil.st.-ni.T-;  iiiv  s,,,,   In  lir  m.M,.;isr.l 
i„    ,,,„,, „,,i„„    I,.   1|„.    .....lilN    i.r   tl..'   .li-'.i-'.    til.'    inrn.iM.    lirin-    i.hiMIv 

,„    ,1,,    |.|;,M,u       Till.    ii,r.,.,M     in    i-l...l.-l.r..l    ,r.,nliu,.l    iiniinly    ti.    tl..' 
|,i;,.„n,       w    ,,;,Hi.-i|l,-.,l>     inhlisli,,...    miiit    ll;-'    >uI..1.,II.m.    i.    IIM,-,t1V.-t  ..,1    ,1, 
;iiii,,iiril   1,\    ,\.-.'-si\i'   li  I'linu   \v  itli   I'ii' 
The  Destination  of  the  Fat  of  the  Blood.     In  ^j.  n. ml.  it  may  !.,•  sai.l 

,|,„  ,|„.  1,1 1  r.it  i.tr;in-|-il-lt.,tl,,v,.  |.|,„....;     1     1  !„■  ,|.-,...t>  I'm-  I al  ;  ''D 

,1,„    I,,,.,.-    iMMJ      ::      111.'    r.^MMs        Til.'    lal     liirM-nl     in    .'iii-h    -r    tlli'S..    lihirrs 

,i„r,',.   fr lint    in   ll !li.'i>.  a^   i^  i-v-l-'l   '■>    -'li'-'i'-^'l   I'N^niiinat .nn 

!,.    , ll,..,U  .l.'^.'l'll"  .1  .  :i    I'li^"  ii<.      Til.'  .'.  /-'    '■■'!   iMlally  >  i.'lils  ;,l...ut 

n.-,  ,,..,•  ,'.',,1  ..1   .1.  mtal  ui'iulil  a.  ra.l>   a.-i.l.     Ti.'  '—"    ./"'•  -n  tin'  ullnv 
1,,„„|     ^i.'liU   i.nh    al...nt    litl    |.ir   -'n'    •■I    i'^    '"'■''    "''-'''    ^'^    '■•'"'^     •"'"'■ 

Tins    .liir.'.'.'li.-i'    iiMrn-.'iI.'>    llial     tin-    lall>     ,'n'i.l    niilM    1„ inlunril    Ml    tl.i' 

M.MU'.    uill,    a    Min.'li    lai-'i-   l,i..l.'.'nK'   Hian    i.   tin'   I'aM'   in    lln'    lal    ..t    tin' 
,|,.,„„,       Tin,   lai'L:.'   nn.l.'.-nl.'   i^   |.M.l.al.!^    tli.it    "i   UriWun    m-   ..lli.'i'   pin- 

,,|„lM.,n.    1    tin'   Miialli','    nn.l.'.-nl.'    ni    tin'    .l.|".t..    ilia!    ■  r    in'Mlral    tat. 

Tl„.  /,,',,    i.'l  laL.'v  an  n.l.'nin'.liai ,'  imMtn.n  |.,'lu,',.n  il.'pot    f.,t   ami  li.sn,- 
1,1    in    it^   vi.'M   I.r   lalU    a.-i.l       Wli.'H   in.  .'n-lis.    .in-taliuliMn   "f   tat    i.   '^n 
,„..  ,„,    ll,,'.  li^.','   fat    1.  Ilk.'  llml   ..I    H,.'  tii-Mi.'^;  Imt    u  I,,',,    lal    m-'tali..!  isln 
I.  ;„.,iw,  tl,.'    Inn-    lal    ...•.■n|in's  an    int.'iiu.'.li.-it.'    l...-itn.n    lii'tw-'i'ii    Iimt 
l;if  ami  .1.  |...)    I'.'it. 


H 


i  <l 


Mi'r 


i 


f 


u 


fi 


m 


^1.'. 


!  n, 


i      '! 


lii 

1  1 
1  1 

-1 

.■     1 

!     i. 

\  {    jj 

7(M( 


Ml  [Mini, ISM 


AiMillit  r  ililTclrlii'c  ill ','  tlir   fats  in   tlir^..  llirrr   |>liii-i'>  is  willl   ivi;:,!,! 

to  111.'  ./.  ./r--  '.'  sill  iii-iil  ill,,  III'  111.'  r,iM>  Ill-ill  lailii'lrs  'I'liis.  it  will  I" 
li'llirliiliiM.'ij,  is  ili.|ii',ili-.|  li.\  till'  inilllii'  \;illlr;  tin'  lii'-'lllT  till'  iiMlili. 
\.ili|.'.  till'  'jn.-i!"  r  till'  i|rs;itiirati.iii  "\  tiiltv  ;ii'ii|.  In  ili'iml  I'lit  tliis  \;ilil 
is  ivliiti\.'l\    l.iw       I'm-  lA.iliiiil.'.  iilii'iit   :;il  ill  III'  '_'iiat   iiiiil  iilmilt   tia  in  tiiaii  ; 

.li'I.cinliiil.'  -M' "lial   nil   llir   I'at   lak.ii   in   tlif   f I,  rniiiji.-iri'il   uilli   \^lii'-l 

it  is  iisitall^  a  liltli-  lii'^iii  r.  Tin'  fat  in  tin'  lisii.'s.  i.n  tlic  ntliiT  lian.l 
lias  a  lii'-'li  imliiii'  \aliii-.  |>..s-il,l\  l|l»  t,,  |:;n,  'i'l'i'  i.iiliiii'  \,-ilil<'  n|"  tlir 
I'al  I.I'  tlir  liv.T  is  iiiiiaikaM\  inr.'iisl.-nit .  Iirini;  almilt  tli.'  smiih'  as  tlia' 
111'  111.-  lissl||■^  wli.'ii  t';ill>  ai-i.l  iintal'iilisiii  is  not  |iartirnlai!\  arliM-.  Init 
a|'|>ii.\iiiiatii"_'  lint  nl'  llu'  ilrimts  wliin  I'at  iiinliili/atinn  is  |ii-.iri'i'.lin'-' 
'I'hr  si.jiiilli'a r  this    I'ai-t    \\v   s|,nll   I'nnsi.l.T  bitiT. 

The  Depot  F.at.  'I'Im'  j.lai'is  in  llir  animal  li.iil,\  w  Ih'Ii'  ilipiil  Tat  is 
.j.'liiisit.'.I  in  ■.M-rat.-t  aiii.ainl  a'r  llir  siil.iMitiiiii'nns  ami  iil  nip  lil  miral 
tissiiis       Tli.'-i'    I'at    .|.'|im|>   may   s..mit!mi's  luiumi'   nl'  rii"i  niniis   si/i-.   a^ 

III     llir     I'asC    nl'    llir     r^llllnl's     .In"     . .  T     I  '  ll  1 1 V .' I',     n  l'     wllnsf     t.'l    ll     I'mh      Wrlulll 

■to  ii.'i-  .'lilt  \' as  .111.'  tn  t'al  r.ln.T  ^irnji"  Is  tlial  111. 'IT  ma',  ii'allv  li'twn 
■  lilTi'i'i'iit  tvji's  .  t'  I'iit  stnia'jr  niu'  nl'  a  imii'h  ti'iiiimi-ai'v  i'liara>'ti'i' 
\'.  Iiii'li  i.'aililv  talis  n|>  ainl  lili'i  at.'-  tlu'  I'at.  Init  \vlii.'ll  1'.  •■'i  !iniit."l 
i'.i|'ai'it\  an. I  pnssililx  niiiliT  tlir  niitml  nl'  -niiir  (jiiirklx  arli'i'j;  n'.'iilaf 
iii'j  iii.'rIianiMii.  liki'  llial  nl'  tli.'  .jh  .-n.j-.'H!.-  fiinrtinii  nl'  tlir  li\i'i-;  aii.l 
an..tlirr  lit'  .-I  mnir  i.rrmanriit  natnir.  intn  w  liirli  tlir  I'.it  is  -.'..wlx  takrn 
il|i.    I. lit    tin'   ra|>a.'it\     nl'   wliirli    is    \rl'\     limrli    'Jfratrr. 

'j'w  n  i|l|i'stinns   |il  I'srllt    till  Ills.  |\  rs  rnlirri'llili'.'  tilis   il,'|int    l':it         1       wll.'lr 

.1,1,',    it    ,'.  nil'    t'lniii.   aiiil    '■_'      ^^llat    1' iii.s   if   it'      l,'r'_'ar,liii'r    "■'     <i,>irri 

1,1   11,1    ,h  fill   fill,  tlu'f.'  is  n,.  . I, '111. I   tliat   il   r'.iiirs  jiaillv   I'mni   tin'   fat   am! 

partlv     finiii    till'   r.ul.nliN.liat"   nf   llr    I' 1  :    in   ntlirr   wnnK.    it    is   rilln  i- 

takrn  via.h   ma.|r  witli  tli.-  I'.i.  .1  ,,\-  inannfarluir,!  in  llir  ..|._'anism.     Tli.al 

snlllr    nf    il    .•nlllr^    finin    tlir    f.lt    nf    f.in.l    is    I1..H     a    Will   .'si  all!  isln'il    f.'lrt.    til,' 

r\  i,li'in-r  fnr  wliirli  iir.'il  iml  ilrlain  Ms  Inn-.'  Tlir  lii-.l  kimw  n  r\ |.  rimriii 
rnlisists  ill  liist  nf  all  siaiviii"  all  aniiiial  iniii!  lis  slnr.'s  ..f  fat  air 
nr;ii-l\  r\liai|str,l  ami  ll'rii  fr,',l'|,'^  !iim  \»itli  s,,i;i,'  ".'ar  iiiaiknl"  fat 
tli.'it  is,  wi'li  s.mir  f,,t  liaviii'j-  a  rliai  arl.iisi  i.'  |>ii,|..rty  wlii.'li  it  will 
lint  Insr  iliirini.'  alis..i'|itinn  It  v  ill  lir  fniiml  llial  llir  i|i  |>.,|  fat  tlnirliv 
il.'Iiiisil.'.I  |.|i'-.'iits  iiiMliv  nf  111.'  ijiialiilis  .,f  tk.'  fill  fat  Tlir  "  ra  i' 
m.'iiUiin.'"'  nf  tlir  I'al  may  kr  srnirr.l  li>  n-ini:  fats  ,.■'  ,liff.'irnt  mrltinir 
|i. lints,  sin-li  as  mntinii  fat.  wliirli  lias  a  lii'jii  M  I*,  m-  nkiv.'  nil,  w  lii.-li  lias 
.■1   ],,\\    M  r      < 'II    f.'.'iliii'..'  a    pi  I'V  i.iiisly   stai-\r.|   .l.r,'   wiili    miilinn    fat,   tlr 

Ml*,    nf    till'    li.'p.il     f;it     .-ippinarlirs    lli;i1     nf    liintlnii     fit       lir    III inrs    a 

il.r,.'  in  slii.p's  .•l.aliin'j  ;  w  lirlr,'>s  '.vli.'ii  ..li\r  nil  is  f..l,  llir  siiliriil  a  nrnlls 
fat  lirrniiirs  .,il\  (  i|-  a',iain  ^^  r  nia\  "rar  m.iik  "  tin'  f.at  li\  rnmliiniii'j-  it 
with    limminr.    w  1  ■     l    tlir    ilrp.isil.'.l     fat     will     likrw  isr    lir    liii  ,m  ill  i/r.I     fat. 


y  V  r   Ml  1  M'.iii  I-  \i 


701 


It   iiiil-l    II"!   !•'■  iniaiiin.'il,  Imv.  ..mt.  i1;;'I    •liiiii'jf  l;il^<'>  Iil:i<'.'  in  IIm> 

l.il    ,|mi,i'_'   il-   ;iliiii|.licili    iihil    iMl'iilr    ll    Ih  inliM  -  ili|"'-il'-'l    ill    til''    li->-ll''S. 

Oil  III iiiiiiiy,  tli«'  ■.t.-imp  .iT  iinlU  i.|iiiilil>    i>  I'lit   lip 'II  llii'   lilt.  fcir.  iis 

we  li;iw  :iliv:i<l>    -'■'ii.  it>  i",|ih,    v.iliir   v  ,r«    I  ■■<■>. Uiv  iilli'idl  iiml   its  llU-lt- 
i,i._r    |„,ih|    rli.-iliuril    illilili'-'    tlir    |.ln.'.'  iil>M.i|'l|nl:,        III    ..llu-r    winds, 

.iltlii.iiuli  till'  iil.vcrli.  .1   lilt   il.M^  nn|  rutin  ly  MiLii.tid  t.i  .•..iif..ilii 

^•ith    llir    '.l(liliiir>     .pi.llith-    -I     ll''    'I'i-    '       •■'I-    V"     it    •'■"'I-'    '"    '••I'lll'-^''    i'l 

'  Ills    llil  IC'tillll. 

■|'li:il  Miliir  111'  llii'  ill  p"l  lilt  1  'IH'"  II'  Ml  (•.■i.l'.li'.'li.it''  is  uill  Uiiouii  In 
,l,H-k  i:ii-..is  WIhii.  I'"f  I  '.;'iii|.lr.  ;  n  .iiiiin.il  i-  l"l  ■  n  l^i'-;''  'jHJiiititifS 
,,l'    (■;ill"'li\iiliil'      :lll.|    I  'I'I     ^>ltllMII    i|('ili'_'    liil|s,-i|l;ir    rMTi'isr.    its    tisslli'S 

I ,,|||<-    |..;i'|.Mi    v.i\h    Till        ll    >    '    ill       "■   stlirl    srirlitilii-    |iriii'f    I'lT   lliis,    we 

,|,,  i,,,|  II, r,!  I"  .,■.!  luill'i  tii.-iii  III'  "I'I  1  s|i  linn  Ills  111'  I, a.  '  ■  •■iii'l  till 
|„.|t.  x.l;",  il  v.ill  I..'  11  I.I.  nil"  11. 1.  sl,,,i;..|  llial  ll'-  I'at  i|"|i-  il'J  in  tin' 
ii-xi|,.>  .'!'  II  i^ii.un  «  I  ii;  i-  ii.rall>  iii  .■  ..^^  "I'  Hi-  li!  H'at  .■..mM  liavi- 
I.. (11  .liiiM-'i  lli'lii  till'  I'at  I'l-  in.'Ii'in  wliii-li  ".'is  in'-aiiwli  \r  iii>tal..'li/,''il 
riic  r\|i''i  ini.'iit  \Mis  |iiil'iirnii  ll  mi  t\  u  ymiir.'  pi'js  l'i-.ini  tlic  ^aiiir  !ill"i- 
.ili'l  ',1'  ap|n.i\iliiMlrl,\  r'|U;il  ui  I'jlit  :  <  lir  v  a-  killi'il  anil  tlif  lAart  iiliiiiUlits 
..r   I'll)    UIkI    llilln'_'l'll    in    till'   1"  ll\    iliti'llllilli'l:    till'   nlllcr    ".IS    I'l'l    Inl-   sl'MTJll 

iMiiiitiis  nil  il  diet   till'  I'at  ami  iimlrin  i-u!;t.-hls  .,1'  wliii-li   ui  r.'  iin-iii  iilcly 

iiscitiiiniil.      Wli.'i'   alter    I'.ui'   nllis   tiii^   iii;.'  Wiis   Killi'.l   iiinl    tin-    tat 

'litrriniiiiil.    it    uas    f nl   tliiit    iiiiu-li    i.m'I'c    Innl    lii'fiMiit'  ilr|)iisitiil    limn 

.'iilllil  111'  inTnlllltiii  fur  l'.\  till'  t'llt  ami  IHntrin  nl'  tin-  t'liml,  rvtii  siipi»ns. 
ill'.'   Illilt    all    tin     iiVililillilr    ciUlinll    of    till'    pint. 'ill    liiid    liiciinic    <'n|iM'ltfiI 

iiitii  fill.     Till'  niily  < Iiisiiiii   is  tliiit   till-  riiilinliyilriitc  must    liavc  lu-i-ii 

,111  iiiipnitiint  siiiiri'i'  111'  llic  I'Ntiii   fat. 
The  Destiniition  of  the  Depot  Fat.    Tin'  drpnt  f.it  lirrniin's  iii..l.ili/..'il 

,iml  triinspnitf.l  li>    tin'  lil 1  to  tlir  ai-li\r  tissiio  wli.iii'M'r  tlic  iiii'f^'y 

n^.plif.Min-nls  <>f  tin'  liiiil.\  i|i'iii;iml  il  |)iiiiii'_'  slar\  at  imi.  as  \\r  Inivi' 
-.'•11.  til.'  .li'piit  lilt  is  11-,, I  t,i  supply  W  |.i  r  i'.  nl  i.f  tlir  .■iirrny  nn  uliirli 
111,'  iininiiil  iiiiiinliiiiis  its  .■\ist,ni-.-.  {'..•I'.K'  tin'  I'iil  is  t  liinspnrtnl.  il  is 
pri.l.iilily  1. till, ,'11  iliiun  intn  fiitt>    iwd  iiml  ul.v  ii'iin.'.  iis  uh'n-li   it    piiss.'S 

i1i|.,,i,,j1i  ll ,.11   \Mills  t,,  l.|,  ipjiiiii    ll  ,"n-liiii'tid   intn   ii.'iitiiil    fiit    in   tin- 

1,1 1.      Wliil    a'^i-iii-v    I'll'.iMs   tlii-   ,-nn  laiil    JTi-iilMlnw 'i    ami    r.sMit  In-sis 

..!'  fill  it  is  diflii-iilt  In  siiy.  Tun  (  si  ,r -pi  it  I  in;:  I'li/vmrs  iiic  prisnit  in 
l.ln.id.  nm-  ii'-tiii'.'  niiiinly  nn  sinipli'  I'strrs,  tlic  ..llirr  mi  ulv  (•.•rid''- ;  Iml 
It  llil-  lic'ii  iiiipi's  ill!.'  tn  d.'ni.in-lrali'  iin>  i  \  .1  nt  i  .  1  it  i"nslii|)  l.i'tu.'.'ii 
''ith.'i'  iif  tli-'iii   iiiid   llic  i-NIi-nt    "I'   fat    iii.,l'ili/iit  ii  ti. 

The  Fat  in  the  Liver.     Tlh'  pli\  si,,|.,.j\   of  tin-  lixct   fiit  Inw  l.c.'ii  very 

ilili..','litly   sti|,|i.'i|,    piirtiriiliirl>     l'\     l.'iitli.'s   iiiid    his    plipiU.         The    niit- 

(■nine  nf  tills  u  nl  k  li.i-  Imn  tn  slmu    lliat   tlii'  IImi iipi.'s  an  i'\t  r.'iin-l.v 

iiitp'.rt.int    p—itii'ii    in   tin'   iiii'tiil"ili-iii   nf   fill,   ln'in'j:.  iis   it    were,   tin'   liiilf- 


! 


I 


: 


11: 


i 


,i 


1 


ii 


'i    \ 


Yicj  Ml  T\f<'l.l>M 

„;n     1 .,■    i„    11,-    |.lr|.n1MtiM||    M|-Il,r    ^.■.tl^     M.M.I    MM.I.MM.!.      f.M     .   .l,„l|l.ll.t  inh 

in    tlf   lisMi.'<,      r.it    is   ii    iMat..n;,l tiui  my    \:u  ■^■-    >iu,u\U\r.   oC    pMl.^h 

lii.l  .iMi-.jv  Wliil.'  ill  ll..'  .I.|-l-  llii^  |"'l.nluil  Mr,.,.>  I,  M.  l."-l<.,|  .luav 
MS  I..  !,.•  u.mv.iil.il.l.'  I.. I  ti->ur  II..V  Tn  iiniK-  I  .r.  ..il.ililr  tl.'  .I.|'"t  l-' 
is  .■iini.-l  Ii.  til.'  livrr.  uli.lr  ill.'  riu-y.-^  1m.-,,„„  ,hI...'K..1  ImH  II. 't  ;i.Mii 
:,lly  lil..M;,l..l      Til.'  r.il   is  tli.'.i  1niiis|..„  l,    I   I.,  tli.     m..ii..,.  ;in.|   lli.'  lil»":i 

llnii   ,,r  111.'  .ii.i-j.N    ."■.•III-      \.iiIimI    Till    i>  III'     ^  .1    ■-'imp.. ■•■. !'•'■:   i'   

t;,iii.   inii.'li    |...t.hti^il   .'ii.'i.jv.   l.ut    n..t    ih    .1    Milt,, I.!.-   Iil'..n    f..r   .'SI'l.' 

s TIm'    liv.r.    IIS    it     U.IV.    .lli-    ll'is    l;,,M|M,^^.l.   .,     Ull.l,.'.'     it     i'^    s.Mt     t.. 

Ij,.'     lisMI.'s     t.l     I \|.l...l.'.| 

Til.'    ■-'r.':il    iii,|..,ll;,iu-.'   ..f   tl,.'    Ii\.i    in    fat    iii.  t.-il.     ,m,i    is   iii.li.'a'.'.l    1.^ 

|.aiis„n  ,,f  ll>,'  |..'n'.i,li, ..-.•>  ..f  fat      .n    Im'II.t  '.f  '  .1-    a. -LI      .•.■uli ! 

ill    ii    nn.l.'r   .lilT,  i.i.l    .■ lili.'Hs    ..l    niiti  iti..ii        In    'l-    ....linan     Mill    .'l 

slM„.jl,|,.r  I s,.  .iniliiaU  tlu'  ll\.T  itaiiis  |-|'..|il  L    L.    »   p.'i-  .-.  nt   ..f  l,i'-'ii'  > 

f;,M\    a.'i.l.   l.ul    ill   al t    ..I,.'   i,,   .•v.'i.v   .•i'-'li'    •>''        ''s  a    ii 1,   lii'jli.'i'   |"'i- 

,.,.,,t:,M,.   V,  ill   I,,'   r i.j   t ,ir      Til.'  sai.i.'  is  1,11.'  in       l...iat.,r>    animals 

1,1    tl,.'    .'as,.    ..r   III.'    Ii,in,an    l,V"i'    as    ..l.lain.'.l    Imni    .    ,l.,i.si.'s    i itain 

..i.iss.s  ..r  (.ati.nts.  r,,„ 1..  TH  |..f  .■.'1,1  ..r  iii.'  ■'■'^   v.i'ji.i "f  th.' 

nv.'ai,,  .,1-  •.':;   |"'i-  .'.III   ,.r  III.'   im.ist    u.'i'.'lit.   iiia>    I'.'    lattv    a.i.l.      Tl  >  i-'    i- 

tli.'i'  ..I'j.in   ill  tl,.'  animal   l....lv    Mial    is  .'M  i-  l..a.|.'.l   uiili    lat    i'     '  ^  >^ 

,'\f.'i,t.      As  ill  III.-  .l.'i...ts.  this  liv.i-  lat   mi'^ht   lii'  .1.  liM'.l  'illn'i-  I'l'"'      •■''' 

,.,,l'ii,-.|   t.,   tl i"jai,    IV Is.'uli.'M'   in   tin'   l.-.lv.   <"    il    mi'Jil    n'|.r nl 

a    si(r|il,|s   III'   niaiinra.'tnr.-'l    lat. 

Tlial    l,„,.^,. "'■'"'■'■"    •■''    '■>'    '"    ""    '""■   •"■'•'"^   '-   "■'>     i''"l'l.^    .Iiun.n 

,,,;il,l,.    Ii,i;l,     in    III.'    Ial".ial.n>     a,;. I    in    tli.'    .'lilii.'.       Al t     fmly    In. ills 

,,rt.r  L'ivin-  i.hl.iil.i'in   t.,  a  .|."-'.   il    I'as  I n   f.'iin.l   tl.al    .'ii.ii'nn.ns  .,ilan 

lili.'s  „f   Tat    .'ipi-'ar  in   tin-  !i\.r,   a    tVu    In, ills  lal.r,  InnMV.i'.   tliis  .'N.'.ss 
,i|-    fat    nia>     liav   .'Utiri'lv    -lisappi-ai  ..I        I'att.v     inlilt  lat  i.ni    .,1'    lln-    li^.i' 

is  als,,   .il.s.iv.il    in    li'ii-plniMis    i...|s„i,l„'.'.    altl uli    in    tliis   .'as,,    ill.'    r.'it 

usually   |,iisists   till    lln'   'l.alli   .1'   tl,.'   animal.      In    man.   in    .l.la'.  .'-!    .'III., 
,.,,l',„i,'i    |M,is,,nin.j    ami    in    .'V.'li.'al    \  .unit  iin.',    .'iininnnis   ,|iiai,t  il  i.  s   ..f    fat 
„ia>    I,.'  i.r.'s.nl   in  tin-  lix.'r  ^' itl'in  ■■<  "■•■'>    -'""'    I"'''""'  "''  ''""'  •'''''■''  "''' 
..Ms.'l    nf   tl,.'    .'..inliti.m.      Tl,.',.     .-an    tl,.  I  .'f.'i'.'    I'.'    m,    .|..iil.t     that     lal     i- 

tians|.,,i't,.l    t..    ll.'    Ii\.'i-    iiml.  1     al.inn  mal    alilnms.    lull    this    .-an    in.t 

!,.•  tak.'ti   as  ,'vii|.',  I-.'   tl    .1   Ih.'    In.i'   h.is  anythim.'   L.  -h.   ^^itll    f;il    ""'''l' 

,,lis,n  in  tl,.'  i....i„al  animal       S,|,'h   .'xi.h'i has  l,.-,.|,   sn|.|>lii'.i   l.y   Cmii..' 

,.|i„|    Mnltram.       u  I-   ha\.'   h.'.n    aM.'   1.,   shuw   that,  al    l.ast    in    I'al.l.its,   a 

similar  invas I   ll,.'  ii^.'i    v.'li   fai  ...-.'n's  in  hit.'  i.iv_'iu '    ami  .ail.v 

l,.|,'larmn       Th.'y   aU,,    f 1    that    ih.'    1.  '^^     a.-i.l    .l.-p.isil.'.l    in    tl,.'    Ii\.i- 

in   lat.'   ,.i.".'na,,'.'>    '^ix.'s  an    ii..|ili.'   vain.'    '•hi.'h    li.'s   In'aivl'  t.,   thai    nf  th'' 
m,-s,-nt.'i'i.'   falt.\    a.'i.l   ihaii   is  Ih.'  .-as,,   in   ni.rnial   animals.      .M.ittiam  <'..n 
.liKJi's   that    ■'"  hi'ii'M'i- 


th.'i.'    is  aliiimlaiit    fat    iin  lal"'li-ni.   th' 


I  \y  Ml  I' Mii'i.i»\i 


-o:; 


liv.T  is  f.iiiiMl  to  l.r  iiililttiil..!  >vitl.  liitH.  |.i.NiiiMiilil>  t'.  Im'  liiin.li'.l  ..n 
..K.'wl.rr.-  \\\u-u  wi.rknl  up."  It  i-  ini.'r.siiti..'  lli.it  tli..  1. 'us  U  irr.r.ly 
111'  iiiisiitili.itiil    t'littv   iii'i'N. 

Tli>'  iiK'st  liki'ly  so'in.  -I  III.  il  lni>.-i„„li-l  I-  'I"  H'lr  U  III.-  I'iit  1»its 
nit  ill  llir  .l.|>"ts.  m.l.s>  wli.ii  .jii-i  »iii>it  i^  i.i  imoul'sh,  «|i.'|i  it  iiiiiv  l.c 
ll„'   I'at    I'iniii   III.'   int. -till.'      Tli.it    niii.'li   '<(  it    .■.uii.'h    fi  hiii   tli.     ./.,.../«   In 

,,i,ily   .In sliMt.-l       Tlni«..   Ih.'   t.i..r.-   .Al.'iisn.'   tli..    iiidlt .  ,iti..n    ..f   tli.' 

livri  uitli  l';it.  111.'  UN. I.'  .•L.s.lv  will  tlu-  fjit  I..'  I"..mi.l  t..  iil'I-.-..  witli  tli'' 
,|r|...|  lilt  ill  its  .'Ii.  i.ii.'.il  .'Iwiiii.t.  liMi.'s  Tliis  liiis  Im.ii  \.iy  ..r.-nrlv 
vIm.uii  I.v.  Inst  ..r  nil.  sljirMii'j  iili  :il;iMliii  s..  as  1..  r\r::v  Hi.'  .l.'|i..ts  ..f  fiit 
;,>  iMihli  a>  |...ssil,|,.-,  IIm'Ii  f.-..liii'jr  it  -111  >"i.i<'  ••.Ml'  111  Ui'>\"  I'al  Mllillsinil 
ni.lliii..' |...iiit    ..r    ;i    l.r.ni  ini/.-.l    fill     :    iiml       II"'    mi'-lli'''    ,liiy    ..r    s,,    ..f 

,,,,i\;ili s,.   ii^   1..  .'I.'.ir   III.'   liv.'f   its.ir   ..I'   I'at.    |M.is.,niii..'   tli.-   iiiilniiil 

uilh  |.I,..s|.|m.ius  ,.r  i,l,l..ilii/ili  Til.'  liv.'i  'ViH  >"•  l''"iii.|  si,.. illy  iil't.'!- 
\viu-.ls  I..  1..-  in\ii.l.'.l  XV  ill.  fill  wlii.-li  liiis  ,ill  111.'  .iiriiiiiiKs  ..f  lliiit  .«" 
« liii'li  till'  imiiii.il  liii.l  li.'.'ii  I'll. 

IM.IrM. f  III.'  siiiiH'  .-liiiriM't.  r  liiis  I... '11   fiinii-li.'.l  in  ii  s.i  i.'s  ..f  cliii 

,,.,,1    .'iis.s    l.y    ,.l.s.l\iiti..lls    .III    111.'    ill.M.llht    ..f    fill     illl.l    'II.'    i..Iili.'    Villi!.'    «< 

ll,,.    f;|ti       ;,,i,|   ..I'   III.-   \i\<v      Tliis    is   sln.iMi    ill    111.'   il. m|'iiii>iii'-'    tiil.l.'. 

I-'m  l^     \.  ll'-  "K  I. IN  I  i: 


1 

1'.    .Ml 

i.iri..    :itntiil;l 

N...i.,mI 
lull..  - 

1 

i.mI  „ 

I'rilll 

lllili.. 

|.h.  i.nioiii  1 
ti<u«    ;.t'.'liii:l 

t.  - 

.  '•'iMIM.'hl' 

lilllV 

,l,:n,;;.' 

s_ 

r..v.i 
\..  1.1 

l-.,l,l'> 

I'lltlll 

.lit 

I'lllil 

1,1. 

l',...h 

ll..  |.ii.iiiM    nil 

Mi.iK.  .1 

111. 
II 

r:il.  1 

I'll,  il  i" 

i:i   ..r   lii.'l  1.  1 

l:ill> 

IJ. 

It. ..II 

l'"l 1 '^1 

.jlilll;;.' 

1.:. 

fl.-.M 

liliv liti" 

1 1. 
1  "l 

.III 

ll.'l  V 

III. .HI  I.   V  \  ll  S 
M  IMS  I'l  I:  .   I  SI' 
III-  l.l:\    «  I  .."  !■ 

IJ   I 

1 '..T 

I  l.r. 

11.1 

V,fi 

IT.-J 

Ul.'i 

•.'.•>.  I 

:i«i. » 
It.o 
\::2 
.".  i.f. 

Illl.!) 


1..IIIM.  \  M  IT 
.11    t  \  11  \     M  ll.-i 

tl.'.x 

11. :> 
n.iii 
ll:...'. 
I.ii..." 
I.i:;..-. 

.m;.o 

!i.-.  » 

SI  11 

ni.l 

s(i  :i 
r.:'..n 
<■.!.,  t 


Tliis  liil.l.'  .'Liiilv  sli..us  tliiit  111.'  lu.'i-.-  f.il  tli.r.'  i^  in  Hi.'  liv.r,  tli.' 
ii.'iir.T  this  fill  ii|.i.i'..i..'li.'s  in  .'liiirii.t- r  tliiit  si..r.'.l  in  tli-'  .l.'|..its. 

Tliiit  s.ini..  .if  III.'  flit  in  til.'  liv.T  iiiiiv  .'.1111.'  .lii.'.'tly  //"."  H"  l"l  '<  ■ 
,.nlh,    „hs.„-h,.l    frnn,    ll.     inl.^lin,     is    iilsn    V  .TV     r.'.nlily    ,l.'ni..iiM  nilil.'. 

Tims,  ^^\wu  .■...•..aimt  ..il  mms  phi.-.-.l  in  iIh'  inl.>tii '  iin.'slli.l  i/.'.l  nn- 

iMiiils,  ill. .11..'  uitli  l.il.'  Milts  iin.l  ulx.'.'iiii.'.  it  m:w  f-"M"l  1'.^  '{'"l"''  '  "'"t 
:;il  pi'i-  .'.'nt  .if  till-  iilis.iilii.l  nil  ai.p.ai.'.i  in  tli.'  liv.'r. 


MICROCOPY    RtSOlUtlON    TEST   CHART 

ANSI  and  ISO  TEST  CHART  No    2 


I.I 

1.25 


<-  ■ 
I- 


132 
|36 


IM 

2.2 


14     IIIIII.6 


A     APPLIED  INA^GE 


.  'IB'  *«.■  -  r^or  -  p^o' 


Wu 

hi 

'    It 

lIMi 


'i  ! 


\i 


't '  !i! 


m|  iit^  -p 


M 


l»     ■    i 


•  i 


^S! 


1 .1 


't  1 


704 


Ml.TM'.ol.lsM 


Till'  .liiii.i.  ini-tii-  I'lMtiirc  of  ii«m:iiiu1  i.il  j>  tint  its  I'Mtly  Mi'i.ls  ;iio  Mil;ilili'  in  stciiin 
.'III'!  :ir.  -MiuiMli  .1,  S..ii.r  III'  th.'  1:1 1 1  y  ;i.iiis  i.l'  I  In-  Iimi-  ;in'  Mihilili'  in  -h  :ini,  I  ml  lli.  > 
:,,v   iii.s;,1iii;ilr.l.      I'.v  .li-iili.ili.iii   in  -■.■:nri  „t  llir   I'Mltv  .'Hi.ls  ul.|:n I   l.y  s:,]i..nirir:it  i.oi 

,t'    ill,.    Inrl.    it    i-    |..i^~lM.'    tcT  ,|,tcllnii,.'    I;,  u    Inll.li    ul'    1  lii'    i'n,n;iillll    "il    lllis    plisscil    t.o    tlir 
li\.  1. 

Siiiiilai-  iTsults  Iiiivc  lu'cii.  iilitMiiu'il  v.lii'ii  misatuiiitcil  t'attv  iiciils,  sucli 
;is  tluisr  i-ciiitaiiii'il  ill  cod-liviT  nil,  arc  f««l.  In  all  tlicsc  cases  llic  rcla 
tiuiisliip  of  tlic  liver  fat  to  t!  at  of  llie  foci'l  is  even  more  evident  lliaii 
that  lietwecii  food  fat  and  di'pot  fat.  I.ci-anse  in  the  liver  tlie  newly  alisorlied 
fat  is  not  diluted  hv  that  de|'osited  it  may  lie  nionths  previously,  as  i- 
the  ease  in   the  eoiineeti\e  tiss'lis, 

'I'lie  iplestioii  now  arises:  ll'//((^  A((/j/«  ».s'  In  lln  fill  ilnrliii/  ih  slini  in 
lilt  lift  I-.'  All  iiidieatinii  of  tlie  nature  of  the  ehanue  is  fiiriiisheil  liy 
oliservinj-  the  iodine  value  of  the  fat.  Tliis.  it  will  In-  remi'inliered,  in- 
dicates the  de<iiic  1(1  whi.li  tlie  fatly  acid  is  uiisatiirati'd.  It  does  Hot 
necessarily  indicate  the  iiiiml  er  cd'  iiiisatiiriited  hoiids  present  in  the  fatty- 
acid  molecule,  for  tl»e  ditVereiice  in  iodine-alisorliiiiu  jtower  may  depend 
not  on  the  iiumlier  of  smdi  lioiids  hut  on  tlie  position  in  tlie  chain  at 
which  a  uiveii  doiililc  I  olid  is  inserted.  Kvcii  with  this  reservation,  liow- 
ever.  it  is  eviih'iit  that  the  increase  oliservcd  in  the  iodine  values  shows 
that  the  liver  has  the  imwer  of  desatiinitinu'  I'at.  The  advantajre  of 
this  ehaii'^re  depends  en  tlM>  fai't  that  tlio  desaturated  fatty  acid  will 
he  more  liable  to  hivak  up  than  the  sitiirated   fatty  acid.     In  other  words, 

the  douMe  linkaL'c  will  weaken  the  chain  with  th nseipience  that  it  is 

Halile  to  fall  apart  at  this  ji'ace;  siieh  at  least  is  th.'  r.atiiral  interi>reta- 
tion  \\hi(di  the  eheniisi  would  put  on  the  result.  It  may  not,  however, 
he  the  correct  interpretation,  for  it  has  hceii  shown  that,  altlimijrli  un- 
saturated fatty  acids  aie  iiioii'  siisceiitilile  to  eheiiiieal  (diaii'_'e  in  the 
lalioratmy  than  saliiiate.l.  yet  when  feci  to  animals  they  appear  to  lie 
iiiiu''  stahle  than  many  saturated  acids.  It  may  then  he  wron<r  to  con- 
cliidi'  that  the  iiit  induct ioii  of  a  d  uilile  liiikau:e  in  fat  n-M-essarily  nipai!- 
the  lialiility  of  the  fatty-acid  chain  to  lireak  at  that  point,  llnwex.r 
this  may  he.  it  s«'eiiis  likely  that  one  function  <d'  the  liver  consists  in 
intro.liiciinr  double  linkauvs  at  jilaees  in  the  falty-acid  idiaiii.  as  a  r.'sult 
(d'  which  the  chain  breaks  at  these  places,  and  tin-  fraiiiiieiits  then  uiulefLTO 
further  oxidation. 

Double  linkaL'cs  may  be  introduced  not  only  in  one  ])lae('  in  a  fatty- 
acid  chain,  but  in  several,  h'or  exanip'o,  it  has  been  fouiiil  in  the  liver 
of  the  lli'.,'.  after  oxidi/iiiir  the  fatty  acids  with  pcrmaiii.'aiiati\  that  oxi'la- 
ticui  products  wri-  obtained  iiidicatini.'  the  existence  of  tmsaturated  acid 
with  tour  double  links,  i'ermaimanale  -in  allvaliiie  soliitioni  i>  used  tor 
detectiii!,'  the  ))ositioii  of  these  double  boll. Is.  because,  wlii^ll  it   is  allowed 


11  ■  ■ !      a  1 
51     I      M 


li 


AT    Ml  IMldllsM 


70;-. 


to  act  oil  iiiisaturalctl  fatly  ai'iils  in  tlic  cold,  it  caiisfs  li.vdroxyl  f,'roups  to 
111'  iiitnxliii'cd  ill  tilt'  |HisitiiiH  ul'  tlu'  (jinililt'  limiiis.  Wlu'ii  tlic  (isiiiatinii  is 
pcrfdrnied  at  a  iiioderatc  tiiiiix'iatiiic,  tlic  fatty  ai-id  falls  ai>ail  at  llio 
liydroxyl  Kr()iii)s.  A  fatty  acid  with  ciulit  liydroxyl  sfrmips  lias  been 
(ilitaincd  in  this  way  finiii  the  livci'  of  the  pijr.     The  iircscncc  of  the  liy- 

liroxyl  gniiliis  has  lircn  cmitii  im-cl  liy  liiidiiiu'  that   ai tdliidiiiidc  is  oli- 

tainod  hy  treatment  with  hrdiiiinc.  An  acid  of  the  smimc  foniuda  is  said  to 
lie  |)res('iif  ill  cdd-livcf  oil.  'Vn  siiiii  up,  we  iinii  cmiclude  that  there  are 
certain  ixisitimis,  in  the  cliaiii-.  (if  carlion  atuins  which  cnnstitiite  tlie  fatty- 
acid  rnilicle.  where  the  liver  intrcidiices  clmilile  lidiids,  and  that  the  \\eal<- 
I'licd  radicles  then  circulate  to  the  tissues,  where  they  lireak  up  at  tliose 
positions. 

Hut  this  is  ]>i'oIiahIy  not  the  only  way  in  whii'h  tlie  liver  assists  in 
the  iiietaliolisiii  of  fat.  It  may  also  take  iiart  in  the  Imildiiif:  of  fatty- 
acid  lailicles  into  the  complex  molecule  of  lecithin.  The  ])roeess  of  de- 
Mitiirafion  that  we  have  .just  considered  is  i)rolialil,\  a  preliminary  step 
to  this  incorporation  of  the  fatt.x-acid  molecule  into  lecithin,  for  it  is 
well  kiiouii  that  lecithin  contains  hiu;hl.\  unsatiirated  fatty-acid  radi- 
cles. Ill  siijijiort  of  such  a  \  iew  it  is  interestiiiL'  to  note  that  in  aleohol- 
cther  extracts  from  normal  and  pathohitric  livers,  the  leeithins.  whieli  are 
lirecipitated  liy  acetone,  liave  hiuher  iodine  values  (  i.  i' .  are  more  lliisat- 
iirated'i  than  the  neutral  fats  extracted  from  the  same  liver,  which  also 
have  liiu'lier  iodine  values  ihan  the  depot  fat  of  the  same  animal.  The 
desaturation  process  must.  ther(>fore.  involve  the  fatty  acids  liefore  these 
lic<'ome  liuilt   into  the  lecithin  molecule. 

The  liver  is  prolialily  not  the  only  place  in  the  animal  liody  wliere  the 
ilesaturntion  of  fjitty  acids  is  Inousjlit  aliout.  The  relative  a<-tivi1y  of 
the  dilTerciil  tissues  in  this  nuard  lias  iicn  studied  liv  feedi!  ^-  eats 
with  fatty  (ish  and  then  determiniuu  the  iodine  value  of  I'at  from  various 
places  in  the  lioily.  The  alisoilied  fat  v\as  more  olivioiis  in  the  liver  than 
ill  the  sulicutaiieous  tissues,  lieeanse  it  had  imt  hccome  diluted  with  fat 
de])osited  it  ma.v  have  heeii  months  previously,  which  would  he  the 
case  ill  the  fat  of  the  lat  depots;  and  it  was  found  that,  althouudi  the 
iodine   value   of   the   suliculaiieoiis    fat    w:is   sli.^htly    raised,    that    of   the 

liver   was   miieli    more   so.    indicaliiiir   that    the   desaturation    pi' ss   liad 

heeii  more  active  in  this  origan.  Iml  had  also  oeciiireil  to  :\  ccvtain  i^xtent 
in  tlio  dejiots. 

Hefore  leavinir  this  suli.ject  of  fat  in  the  liver,  it  is  iiiiportant  to  re- 
call   the   old    oliservation    of    Ifosenlhiil.    that    a    mor ■    less    recipr<ical 

relationship  exists  lietvveeii   silycoiien   nd    I'at   in  the   liver.      When   much 
irlycopen  is  present  there  is  little  or     o  fat.  and  vifi    n  rfn      It  is  impor- 


i    i 


!,  i» 


I , 


1 


£i     *H 


:ii. 


i 


7(111 


Ml   I.Mlol.IsM 


\]W' 


tiiiil  t..  iiiiti-  IliMt  llif  cxncl   l.iciilicms  .if  fat  nil. I  (•iirl»..li.V(lriito  in  llic  1h 
])iitic  l.iliiili'  iiic  soiiicwliat  (litTci-.Mit  in  tlic  t\v.)  casos. 

A  i»ra.-1ical  rliniml  iipiiliciilioii  of  the  aliovc  work  is  r.mii.l  in  tlio  fad 
tliat  fats  will  he  nnno  iratlily  iilili/.c.l  by  tli.'  li.i.ly  when  tliey  cimtaiii  .1 
liisili  ]>.M'.'i'ntai,'.'  ..f  iiMsatnrattMl  fatty  aci.ls.  It  is  pr.iliably  fm-  llii- 
icMs.in  tliat  Norwo-riaii  ci.!  liver  oil  is  of  such  tiii.hnihted  nutritive  vain.' 
It  is  luii.'h  more  so  than  XewfountUaiitl  .■o.l-liver  oil,  he.'auso  in  the  jir.-p 
aration  of  this  variety  oxidali.m  oeeurs,  whieli  makes  it  n.)  longer  uiisat 
urate.l.  Fish  oils  in  fieiieral  are  more  unsaturated  than  other  aiiimiil 
oils,  and  are  for  this  reason  more  nutriti.ius. 

The  fnt  in  fin   lis.tucx  difi'ers  very  materially  from  that  of  the  liver  .n 

the  depots,     duly  tiU  per  eeiit  .>f  this  fat  isists  of  fatty    -eid,  wliieh  i^ 

i)iTseiit  very  largely  as  i>art  of  the  lecithin  ni.ilecnl.'.  thus  acc.mntinR  foi 
the  lii'-'h  io.liiie  value.  Some  is  i)rolialily  als..  present  as  simi)le  u'lyeeri.h  . 
in  a  hii-'hlv  unsalural.'d  ami  tli.'rcf.ir.-  very  fragile  cr.ii.liti.m. 


!         i 


Jl,\ 


(Haiti; I.'  i.xxix 

FAT  MITAr.oMsM   C.ntMi 

Twii  viTv  iiji|iiirliiiit  i|uc'>linii>  n|'  tatty  Ill-id  iiiftalKili-.iii  may  now  In' 
(•(iiis'ulcrcil:  namely,  il^  limi-  is  falhi  miij  fnniKiJ  fram  r-irl)i)l(  i/ilrah .' 
and  (21  ifhnl  lucniiux  of  lln  friniiiii  ills  Into  irhlcli  tlir  f<ilt>i-iirlil  molrciih 
in  split  lis  till'  nsillt  nf  till  ill  siifiirntinii  jn'iiri  s.<.'  AltlloUftll  flicsc  prol)- 
Icms  involvo  clicmii'al  details  <>(  a  somewhat  eoni])lex  nature.  w(>  must 
not  on  lliis  aceoinit  fail  to  eoiisicjer  lliem;  for.  as  we  sliall  see,  niiieli  of 
what  is  known  has  an  im|ioi'taiit  piaetieal  applieation  i|i'peHd'n<i  i^n  the 
I'aet  tlint  eertain  of  the  inteiiiieiiiai\  snlistanees  nia>  aeeumnlafe  in  the 
iii'iranism  .md  <leveliii  a  toxii   action. 

The  Production  of  Fatty  Acid  out  of  Carbohydrate,  h  w,-  phno  tin 
I'oiiimlas    foi'   f;hieose   an-l    paliiiitie    acid   side   l.y   sidi'.    (hus: 

CirOH-  (CIIOlIl.    Clio    i;;lll."Mi,  Mll.l 
ril,- (riT.I,,-  fY)(Mr    fi.Mliiiiti.-  :iciil): 

ue  shall  see  that  this  tiaiist'oiniatiun  iinist  iiixolve:  t\)  a  coiisidel'alile 
alteration  in  the  stiuctiirc  of  the  molecule,  (2i  the  :emo\;d  of  oxygen, 
iind  i^ii  the  fusion  of  several  <_'lucose  Uiolecules  into  one  mnlec-ule  of  t'atty 
acid. 

The  eonvotsion  of  carhohydiate  to  fat  iheret'oic  in\olves  a  process  of 
leductioM.  and  the  resulting  molecule  must  he  eai)al)le  of  yielding  inore 
energy  when  it  is  oxidized  than  the  oriirinal  one  of  carhohyilrate,  for 
ohviousl.v  the  system  (>_,  f"II,  fMhich  corres|)onds  to  faf*  will  develoj) 
more  energy  than  that  of  <V.  (TI(^  iWhieh  cori'cspond.s  to  carlioh.vdrate'^  ; 
.just  as  w  piece  of  wooil  when  it  is  burned  w  ill  develop  inori'  heat  than  a 
j.ieee  of  charcoal.  This  explains  why  one  i^ram  of  fat  yields  ().:?  ealoi'ies 
of  heat,  and  one  gi-an  of  cariiohydrate,  only.  4.1  I  page  ,">;>.")).  Fatty 
.icid  thei'efore  contains  more  jiotential  energy  than  suirar,  and  in  explain- 
ing its  synthesis  from  suirar  in  the  animal  liody  we  must  indientc  fhr 
siiiirrr  of  tin  i.rlrii  i  iiiri/ii.  This  is  dependent  on  o\iilatinn  of  some  sugar 
molecules  which  do  not  theiiiseh cs  liecome  chanired  to  t'atty  a<'id  - 
proceeiling  siilc  1).\  >iili'  «itli  the  n-duction  uliich  alVccts  the  others  and 
rei)rescnte(l  in  the  outcome  of  the  icaciioii  liy  the  comliustion  jjroducts 
CO,  an.l  If/),  thus: 

IUVH,,0„  •  l.>  (»,     2n  ('(I.     (',,,1I.,.,(_\.  •  'jri  IFd. 
( irlucnse)  I  fatty  acid  i 

707 


•:t    ■     ■ 


TOH 


MKTAlinl.ISM 


t',  I  r^  i 


II 1 .1 


iH: 


! 

1 

'm 

Pi    •      i 

1 

:, 

'!•  • 

liHI 

hUii',ii|!i.i 


i^ 


^'1 

It 


t .  't 


'Mti 


i      'i 


ill 


il 


I  &n 


1:1    ! 


Wlial  cv'kI.'Ih-.'  liii\f  \\<-  thiit  ^iiili  II  /</■"(■«  vv  iiiliiiillii  iirrtirs  in  the  boihi.' 

If  \vi-  coinpiirc  till'  iiitiilif  (if  MNyacn   willi  tl utpilt   "l"  carl)"!!  (lioxidc 

III  the  iTsiiiri'il  air,  wr  shall  tind  lliat  usually  tlific  is  less  of  llic  lattoi'; 
llial  is  1(1  say.  tlic  rcs|iirat(ir\  (|U(iticiit,  as  this  ratid  is  called,  is  us\iall\ 
less  tiiali  unity.     Diiiiiik'  the  f\li'iisi\  c  cinn  .■isidn   dt'  .■artmliydralc   iiitu 

lat,  linwfvcr.  wliicli  ( iirs  diii-iii','  ttic  tall  iii(iiitli>  in  liiliffiiatiim:  aiiiiiiaK. 

:li('  I{.(^  has  liccii  rdiind  td  lisc  as  liisih  a^  1.4.  The  'jrcat  cxc'ss  di' 
CO.  diilimt  dvcr  (>  intake  "hi,-h  siieh  a  (|ndtieiit  indicates  cdiiJ'dniiN 
w  illi   the  nlid\  ('  ci|ua1idii. 

The  entire  dis>iinilarity   in   cheniical   strncliire   hetuecn   the   iiii)lecules 
(.''  fat  and  carlidhydiate  siiL'iresis  that  the  iiriiiiary  step  in  the  cdiivcrsidii 

must  lie  a  thdrduuh  lireakddwn  i>\'  the  eailidhydratc  chain  into  ( ipaia 

ti\-ely   simple   iiidleciilcs.    fnim    which    the    fat    iiidlcculcs   are   then    rccnn 
stillcteil  and    the   Uinieee>sai\    d\\  lien    set    free.      The   pnililelll    is   to   ascer 
tain   the  chiiiiieal   structure  dl'  these  >implci'  mdleciiles  and   th(>   manner 
(if  their  unidii  iiitu  fatt\    acid. 

(  n'  ill.'  -  \ I   -ii;;l;i  »li..ii>.  wliiili   li;i\c  Ihimi   mi;i.|i-.  i1i;iI   ..I    Snii'.ilcv "  ■  m  .■m-  llic  nn.- 

liki'ly.     .\-<  Mill  I'c  -icii   riiiiii  till'   r.'iinwin^'  ,dii;ilii.iis  tin'  lii>t   >tc|i  is  llic  cniuTrsioii  nl' 

t,'liii-c.sr   tl.   |.viiui.-   ;iiiil      \>:v^<-    i\'"'.    Nn.    I    i |ii.iti.iiiv  > .      I!y   cii/viiii.-   aclii.n    ]iyiiivi.' 

;,,.i,|    |.  ..,irn.  i>,.,|    iiil.i  :i.-rt;.i.|cli\.li'      N".   -        wl.icli   tlicii   r,.|iili-ri>(-i   uill tin  I    |.M-n\i. 

;„.|,|   ,„.,|,-.nlc  Im   I..M1,   :i   lii^hiT   krlmn.'  ii.'i.l    (X-.  ■"  i .    riniii   wliicli   l.y  tlic  l.iss  of   CO,. 
;is  ill  till'  c.-isc  i.r  till'  pi.i.li.lii.ii  ..f  ;iiitiil.lrii>.lc   ir.iiii  |.yiieii'  ai-id,  .-in  :ililcliy(lc  is  pi" 

,|i 1  ,  X,..  1 1.     Tills  liiulici  iil.lrli.Mie  l.iliMvi-.  like  :iiTt;iI.l(liyile  ill  !it;nin  ninil.inint;  witli 

[lyiiivic  11. -i. I.   riiniiiiiu  ;i  >lill   liiutii'i    kct.  iii.'  m-i'l  :   :i".I  >"  "n  until   :it   ki-t    ;i  l..iiLr   fiiti.^ 
n.'i.l  cliiiiii  is  l.iiilt  iiji.  tliiis: 

(  1  I    <'.1I,0.,  ^  ()    — lU'llfOCOOJI   ■  I'M  <) 

.;  m|i„.,,sci  (  |.\  ii;\i.'  ill'!-!  .' 

(L'«    CII  COCOOII        <l!  <|tM  .  (It 
;i.vt;il,Ii'li.Mlc 

(.; .  (11  (iiii    (I!  rci  (1(111 .— ni  ciiMin'dcooin  ii.o 

(iiiis:itui,-itcil  kctiinic  iieiil) 
r  ( 1   CII  (11  :  (III '(>('(  Mill  =;('ll  CIIMIU'IIO  >CO. ;  ami  ."(i  mi. 
(liiL;li.T  aMcliy.lc) 
(T))  ri.im  till-  kct.iiiii-  iil.lrliy.lc  t'lHincl  iit  :iii\    sta^c,  an  uiisatinaleil  fatty  aci.l   («illi 
niie  less  Cat.iiii     i-  iia.lily  I'.nMir.l.  lu.il  iliis  l.y  lakini;  up  11  may  l.cedinc  saturati^.l : 
(11  <ll:(  II  CO  CdOII     :  O-  CIL  CII:C||  coon  I  CO.. 

Dinin-  llic  l.iityii.   .-i,  i.l  fci  iiinita' i   u  ■<(  ^w^-.kv  a  slij;lilly  .litVereiit  pn.ici-s  may  dcciir 
iiaiii.-lv.  til.'  laiti.'  :i.  i.l.  ^Iiidi  w.'  kin.w  is  i,. a. lily  pi.i.lu.  ...I  Imuh  -Inc. .so,  may  lireak  (Imvii 

iiii. ftal.I.hx.I''    '  iiiiil  f'Uiiii''  a.i.li,  an. I  tun  >ucli  iii.ili'iulcs  (■.■inli'iisc  tn   fmni  ,'Jii\.v- 

l.utM-ii-  al.lcl  s.!.-;  1  tliis  iii^aiii  i  ..ii.lci:sc  m  fi.rm  Iii-licv  fatt.v    a.i.l>.  <U<>-: 

ii)  ciro,  —  ;;cii,cnoiiconir, 

('uliKMse  1  ( lactic  acid) 

C'.    JCll  Cll()llCOOII  =  L'CII,CIIO  J  ll.COOM 

I  acctaMi'liyili'l 
,:;.    L'ClICIK)    -CiiCIlOllCll.cilO;   an.l   s..  .111. 
1  ,i  iixyliiityiic  alilcliyilc') 


FAT    MKT.MlOI.ISM 


70!) 


That  liiKlMi-  fatly  a.'i.ls.  sii.Mi  •(>  cainoic  (<'II,/';l  '■'"'  •■!'l'r>lic  'f',lf,,0,),  liavo 
a.lMallv  l.vcii  isolatc'l  rri.ra  tin-  |.in.|iii'l-.  nf  ll,i^-  rrTni.'iitafi..n  is  a  very  ^i^nilicaiit  fart, 
ami  it  i«  of  iriliicst  ti>  ii"ti'  tliat  Lcatlics  hut  sonu'tiiiics  fuiiinl  aa  ia.Ti'aso  in  lii^lii'i-  fatl> 
:i,r.ls  t"  I.,', Mil  .luiin-  \\u-  asr|.lir  iaiaiLatl.  n  of  \'i\>y  i.ul|.,  Tafm  tanatrlv.  linwcvcr.  tla- 
ianva-r  c.f  fatlv  a.i.l  r.iuM  not  l.r  v1i,„m,  (,,  ]„■  ntV.-.l'-.l  !■>  .i.l.linu'  -ut'^^taiaTS  I.,  llic 
liviM    ulii.-Ir.  .■i.r.M.lia'j  t..  tlir  al...\.'  ri|irili..n<.  >Ih.iiM   v  irl.l   ri.Hy  a. a. I. 

The  Method  by  Which  the  Patty  Acid  is  Broken  Down.  In  tli<  <liiiiii 
cal  l;i!inr;it(ii-,v.  onliii:ify  oxiiliziiiL'  jiiivnis  attin-k  ll'f  I'jitly-iicid  diiiiii  at  tli.' 
('•;it<iiii  lii'xt  till'  <-ailiii\yl  (('(idlli  sria.up  itlic  ;ii|>li;i  ( 'jitniii'l.  Hill 
this  c.-ni  iHit  (iia'iir  in  tin'  iiniiiial  Imdy.  Iiocjiiisc  it  would  li-iivc  liidiiiid  .'i 
siiinllcr  I'liaiii  cipiitjiiniiiL.'  iiii  iiiicm'ii  iiiiiiilaM-  nt'  ('-atoms,  and  siu'li  cliaiiis 
aia-  never  fmind  iireseiit  in  the  animal  fats.  <>ii  the  ediitrnry.  tlie  cuiii 
iiiiincr  fats  all  cdntaiii  an  even  niimlier  <<(  ('  atuni^  tiins;  I'lity  i'i<'.  <'|IU(V  : 
pallllitie.  (',.lf  :<>,:  stealie,  C.JI  ,<»,:  nleie.  (',,1!    ,<>,. 

The  iiileiiiH'diary  snlistaiiees  wliii-h  aic  priMluced  iliiriiiir  the  y:radiia1 
lucakdiiwn  (if  the  I'alty-aeid  iin.leenle  in  the  niniiial  animal  are  of  a  very 
tfaiisitdi'v  ehai-aeter  so  much  sn  iiidee.j  that  it  is  impossihle  for  any  one 
of  them  to  aeeiiiiiiilale  in  siil'lieient  amount  to  jiei'mit  of  isolntioii,  or  even 
deleetioii.  Iiy  (diemieal  means.  How  then  are  we  to  iih  iiliffi  ihr  inlrr 
innlinrii  pntdidt^'  This  ha-  heen  rendereil  possilile  liy  the  discovery  that, 
when  anythinj.'  oeeiivs  to  distnrli  the  normal  eoiirse  of  fat  iiu'tabclism.  as. 
for  example,  when  the  tissues  are  deprived  ol"  earliohydrates  ''as  in  star- 
vation or  ill  severe  dialietes  ,  the  oxidation  of  the  fatty-aeid  ehain  stoiis 
short  when  a  ehain  of  four  ('atoms  still  remains  iinlirolien.  These  last 
four  f'-atoms  seem  to  form  a  residue  that  is  more  I'esistaiit  to  oxidation 
than  the  remainder  of  the  fatty-aeid  ni'  vule.  It  is  a  residue,  tlierofon'. 
which  is  (piite  icadily  fiii'ther  oxidized  to  CO,  and  IF./)  under  normal  eon 
ditious.  hut  wliich.  althoiitrh  incapahle  of  liecominir  i-onipletely  oxidized 
when  the  mi-taliolism  is  upset,  does  undeiiro  a  partial  oxidation,  result 
iu','-  ill  the  ]n'odnctioii  of  various  intermediary  iiroducts.  These  accumn 
late  in  the  body  in  suflieient  amount  to  o\erfIo\\  into  the  urine,  from 
which  they  can  lie  isolated  and  identified. 

The  fatty  acid  with  4  ('atoms  is  hutiirlr.  ( '11  CHJ'II.rOOlI,  and  the 
first  oxidation  ))ro(|uct  formed  from  it  in  the  hody  seems  to  be  f)-n.nihiil n 
rir  drill.  ( 'H  (  IK  »I|(  11  .('<>(  »ll.  This  then  liccomes  oxidized  to  form  a 
i.ody  haviiii.'  the  foi-mula  ( 'II  < '( »( 'IIJ '(^OH.  iin  tonci  lir  nrliJ.  which,  mi 
further  oxidation,  nadily  yields  Cil  COCII  .  or  untonr.  Tiie.se  siih 
stances  (/i  iixyliutyric  acid,  acetoacetic  acid  and  aeetone;>  appear  in  the 
urine  during  earliohydrate  starvation,  as  in  diabetes. 

It  miKlit  be  oli.jected,  however,  that  a  cliemieal  process  (U'curriiiir  iindei 
.ilinormnl  conditions  need  not  also  occur  in  the  nnniinl  animal.  That  i' 
nrohahlv  does,  howe\er.  is  indicated  by    tlie   results  of  the  cxpeiimen'- 


i 

I 


-I, 
I 


II 


in 


li 


pr 

r, 

1;1 

!; 

'  I 


ill  rl^ 


710 


Mn'AllDl.ISM 


iif  Kiic'Kp  Mini  I'l'  MmiImIi'm  mill  lii>  iMivMirki'is.  Kiiimi|i  ciiiici'iM'd  tlu!  iili'.i 
ipf  iiittiiclnciiiy  iiilii  till'  t'attx -iiriil  iiinli'ciilc  miiiic  uiiiii|)  wliifti  is  icsistiiiil 
to  (i\i(l.ilioii  ill  till'  IhmIv.  Till'  plu'iivl  L'l'Kiip  iC.ll  >  was  t'lUiiiil  to  Ikim 
this  I'lTi'i-t  My  fc'i'iliiiu  ;in  iiniiiiii!  with  thi'  |)h('iivl  (li'ii\  iiti\  t-s  of  iici'tii'. 
|ini|iiiinii',  hiitviii-.  iiml  vali'iii'  ai-iils.  it  was  I'liiiinl  that  the  mini'  cnii 
taiiiiil  I'ithiT  lii|i|iiirii'  sec  |i:i;;i'  ll'lH'  or  |iln'ii:ii'i'l  lli'ii'  :ii'iil.  I'mtli  nl 
ihi'si'  iilf  i'nlM|Hili;nls  of  iiloirialii-  iiriils  with  ul,\  I'ocoll  or  ,illlilloai-il  ii- 
aciil  iCIl  Ml  (  ■<  •<  »||  I,  oiii'  ol'  till'  inott'ili  Itllildilisistoni's  Mini  al«a,v- 
;i\,lil,il'li'  ill  till'  or'_';illisiri    to   I'onil   ^llill   ro|ll|iol|llils.   llllls; 

(1      I'll  ((Kill      (11  Nil  COdll  -~('.IIV<'MI''H,   COOII. 

(  l.iMi/.'h'  I  :;1vKM-..iri  ( lii|i|iiiil<'  iiciil^ 

ii.'i.ll 

iDi  ('iiciicooii    ('ilMi.rixill      (',11  (11  (OMiril/'ooil. 

(  |.lirh.\l;ii.'li.'  i'::lvi'iH'.,ll  .  f  i.liiiiai-rtiirii'  iii'i.l  1 

Whiii  citliiT  licii/oic  ai'iil  iC,  II  '•  "•111  or  phciix  lacctic  aciil  ( ( '„lf  .* '"  ■- 
('(Kill  I  is  I'oiiiicd  ill  tho  I)0(1\  as  a  result  of  tin-  oNidation  of  plionvl 
di'iivativcs  of  the  hi-ihci'  fatty  aciils.  Ilic  acid  coiiiliiiics  with  i;lycocoll 
jH'coidiiiu  to  fill"  aIiov(^  ('(luatioiis,  Kroiii  this  it  follows  that  if  oxidation 
oci-iiis  so  that  t\M>  Catoiiis  ait-  thrown  otV  at  a  time  i /?-osidatioin,  fatty 
acids  with  an  even  ('-atom  chai;.  'loiild  \  icld  hippiiric  acid,  and  those 
with  an  uneven  chain,  phcnacctiiiic  This  was  loiiiiil  to  he  the  case,  as 
till'  aci'oiiiiiaiiyinu-  talilc  shows. 


iiXI|i\l  lov 

rKohiTT 


KXCllKTUl  .\S 


l!iii/"i,-   a.i.l,   (•  II  .COOII 

I'll.  iivl:,.-.  tir    .ni.].    C.II.CII   .COOII 


Vnt   ci\i.li/ri|  lli|.|ilirir  liclil 

N..|  n\i.li/..|  rii.-ii;iii'liirii' 

i.-i.l 

l'liiri\li  H.|.i.iiii,-   a.  i.l.   I'll   .III. CM. .COOII  c,ll  .COOII  lli|.|Mnii' iii'l.l 

I'll,  iull.nl  Mi.'  aii.l.  (■  II  .Cll  .("ir.C'H.rOOn  ('II    III  .COOII  I'li.'tiaiclMiii' 

a.-i.l 
I'lii'iixhal.rii'  ail.l.  f;H,,.rir,.CH,.Cir.CII,.rOOn       (•,,ir,.<'()OH  IlippinJiailil 


fKiiiiii   I)akin.> 


Kiiilidcii"s  cxpi  riiiiiiils  arc  cipially  convincini;.  lie  studied  the  forma- 
tion of  acetone  in  delihiinated  Mood  peifiised  Ihi'oiiirh  the  freshly  excised 
!i\er.  Xoniiaily  only  a  t  race  of  this  siihstancc  is  lornicd.  hut  when  fatly 
ai'ids  with  an  excii  mmilier  of  carhoii  atoms  were  addeil  to  the  lilood, 
they  gave  rise  to  a  mai'ked  increase  in  acetone,  whereas  those  with  an 
uneven  eliain  failed  to  cause  any  ehansre.  The  neetone  was  "ound  to  he 
derived  iniiiiediatcly  from  acetoai'dic  acid.  The  follow  ins,'  talile  shows 
ihe   results. 


KAT    Ml.TAIiOI.lSM 


711 


yOItM.M.   KATTY    A(ll> 


KiiKMATIIlN  OK 
.Ml  TDACKTIr  A.  ID 


\,;<W  n.'i'l  ClI   .CIMIII 

|-r,.|,i,,i,i,.   :i.i,1  CIl  .ni  .rodll 

lintvti,'  .iri.l  (•Il,.<'ll  .(11  .•■(Mill 

Viiliiic  iicici  ril  .ni  ,(II  .<||  .COdll 

M.i.l  CM   .CIl  .<1|   .(11   .(11   .((1(111 


riipriiii'  ;i'  i.l  (  M   .('11  .<  II   .(II  .'  II  .>  >"'ii 

||,,,txlir  ii.i.l  CIl   .CM  .(11  .CM  .CM  .ell   .('(lOII 

(),t..i.'  ri.i.l  CIl   .CIl. .CM. .(11  .CIl  .CM   -CIl  .(  (lOll 

N,„i,         i,i,l  Cir.Cir.(  ir.CII  .CIl  .CM   .C||..CII,.C()(>II 

I), ,„.i,i      CIl  .cii:.cii:.cii  ■cii:.ciL.cii,.cii,.cijL.(  M..c(i('M         

-         ~~  "  ( Kroin    |):iki?i.> 

For  a  Idiii:  tiiiio  it  was  (linicull  fm-  clicinisls  t.i  iiiidcistimd  liow  sneli 
M  pn.ccss  of  nxidation  at  the  /iCalnm  roiild  cccur,  since  tlicy  \v('ri> 
iinahic  to  l>iin-  it   alioiit   in  tii''  lahoiaton    liy   the  us.-  of  tlic  ordinary 

oxidiziiiu'  aiicnts.  Imt  r iilly  Dakiii  liiis  iviiiovcd  tiic  diflicidty  i>y  sliow- 

iii-r  that  1i\dro>;cn  i>iro\idc  ill,<>,'   "xidizcs  fatty  acid.s  jdst   fxactly  in 

tliis  \va.\'. 

We  may  sum  up  flic  results  of  tlieso  e\|)eriiiieiits  and  oliservatioiis  liy 
statin-.'  tliat  imninil  sill 'irati  <l  fnltn  iiii<h  mid  tin  ir  i,ln  inil  ihr!r<iliris  cm: 
mKlrn/o  oj-Ul-^'nii.  iKil  oiihi  in  Ih,  miiiiivl  Innbi.  hut  nho  in  ritro.  In  such 
,1  nninnrr  thai  Ih,  tico  (or  sonii  mnltiiAi  thcnof<  tirniini  Ciilnmx  an 
rnnorril  at   (dch   sncci  ssiri    .v^ /)    in    thiir  <li  cuminisilion. 

Hut  wo  must  not  lie  too  iiasty  in  concludiiiv  from  these  experiments  that 
the  steps  in  tlie  process  are  m  <  i  ssnrilil  in  tlie  order  of  lirst.  the  iirodue- 
tioii  of  a  /Miydroxy  aeid,  an. I  se.-ond.  the  oxidation  of  tliis  to  a  l<et()ne 
._'roup.  The  mere  preseiK'c.  si.h'  hy  si.h'.  of  /Miydr.)xyiiutyri.-  acid  and  of 
acetone  in  tlie  above  experiments  does  not  indicate  which  is  the  ante- 
cedent of  the  other,  and  indeed  theic  arc  several  experimental  facts  that 
seem  to  show  that  the  hydroxy  aeid  may  l>e  derived  from  the  l<etone. 
For   example,    when    accfoacetie    acid    is    added    to   iiiim-ed    liver    and    the 


■'or   example,    when    acetoacttie    acid    is    a 
mixtiiro  ineiihated.  /i-hydrox.\Jiut\ric  acid  is  form 

althoiiirh   lc>s  usually   the    i-evcrse   action    idxidationi    may   ur   when 

/Miydroxy  acid  is  adtled.     A  reveisililc  reaction  must  therefore  he  c 
of  (leeurrinsi-  hetwcell   these   1u(i  sulistalices,   thus: 


i(-<l   I  a  reduction  process 
whe 
he  capaMc 


iiclni'tidii 
CM   .CIl()II.CIl,.CO()ll    < — 

.■v'hlriti   11 
I ,;  owliUtviii'  -.ir'u] 


11  .Cd.C!!  .COOII. 

i'!l,,ln|,.-,'ti.'   !l.-i.|   , 


We  know  practically  nothing  as  to  the  conditions  dcteriiiiniuK  wliether 
oxidation  or  ivduetinii  shall  predominate,  hut  there  are  two  siirnitieant 
facts  that  one  should  hear  in  mind:  (U  that  a  iileiitifut  supply  of  <ixy- 
licu  is  necessary  for  the  oxidativ.'  process,  atid  CI)  that  the  iireseiiee  of 
readilv  oxidizahle  material  in  the  liver  (e.jr.,  earhohydrales^   may  d(tor- 

.1  1*  .•  __!'     I.      <1.         *;...i     .1ir.  II     4. 1  I.-..  l4      iL'     i>i>«kitii(tii)\-     vniil 


mine  the  dire<-tion  whiidi  the  reaction  shall  take.     It   is  commonly  san 
that  fats  hum  in  the  liic  of  carhohydrati's.  and  it   may  he  that  the  un 


i  Ml 


M: 


h 


t    J 


.1 


In 


I  . 


5.1. 


('! 


i      > 


; 

■  1         ; 

-1    ; 

if 

1;: 

ij 

i'  '^ 

\ 

\\ 

712  MK.T.\W»I.I8M 

tloiiliff-l  iliiiiiiiiitiiiii  ill  iiriili'sis  uliicli  (ii'iMiis  ill  dialx'tcs  wlicii  fiirb"- 
li\ (Irate  I'ihhI  is  mivfii  is  (lf|ii'iiiliiit  upon  flu-  dirci'tivc  iiiMiii'iice  wliirli  its 
■■(.iiil.iistiiiii  ill  till'  liver  lias  oil  the  iilioNe  prneessos.  Hut  \vc  imist  !"• 
einitidiis  imt  to  tidiisler  results  (.litiiiiied  Iiy  experiments  "itli  iniiieed 
liver  ill  .juilu'iiii.'  <>l'  tlie  reiietioiis  wliieh  oeelir  iliiriiii:  life,  riovisioiiiillv, 
then.  \\i'  iinist  iisMiiiic  either  that  /Miydnixyliutyrie  aciil  i«  a  iiivessar\ 
xtaue  ill  the  o\iiliiti(>ii  of  liiityrie  aeiil  or  that  it  is  formed  l>y  rcduetion 
of  aeeloaeetie  aeid.  wliieli  is  really  the  lirst  step  in  that  proeess. 

Of  eoiirse  there  is  lio  e\  ideliee  iti  the  aliove  experimeiitH  that  the  hijjhei 
fatt\  aeids  are  also  hrokeii  down  Ity  the  removal  of  two  ('atoms  at  ii 
lime,  nor  has  it  heeii  possilile  to  deteet  any  keloiiie  or  /Miydroxy  deriv 
atives  of  Iheiii  in  the  animal  hoily.  We  eaii  only  reason  from  analo'.'v 
that  a  similar  process  ma.v  oeeiir.  althou'.'h  some  sup|>oit  is  furnished 
for  such  a  \  iew  li.\  the  faet  that  ketonie  t'alty  acids  have  lieen  found  in 
\  ejretalile  iu'i;anisms. 

What,  then,  it  may  he  asked,  is  the  relation  of  the  desatiiration  of  fatty 

acids  which  wv  have  sci curs  in  tin'  liver  (and  prohalily  elsewhere"!  to 

the  H  oxidaticni.'  There  can  he  no  ihrnht  that  liotli  iiroeesses  can  occur 
in  the  animal  liody,  indeed  in  the  same  (Huaii,  e.i:..  the  liver;  and  it  is 
imporlant  to  ascertain  their  relationshi])  to  caidi  other.  The  conclusion 
which  would  seem  to  conform  liesf  with  the  known  facts  is  that  the 
desatiiration  piocess  occurs  (in  the  liver i  so  as  to  hroak  up  the  Ions: 
fatty-acid  chain  into  smaller  chains,  wliicli  arc  then  caiiahle  of  fi  o\ida 
tion  fin  the  tissui-s  ;  desatiiration  may  he  the  process  liy  which  the  mole 
ciilo  is  romrli  hewn,  and  fi  oxidation  that  hy  whicli  the  resultiii!;  pieces 
are  fiiiallx  split  to  their  smallest  pieci's-  that  is,  to  molecules  of  the  si/e 
of  acetic  acid,   wliii-h   mi    fiiiitlhi  completely   hiirnt    to  cailionic   acid   and 

watei'. 

Tlic  iiii'iiMM'  I  r  icMllui'  \:ilui'  cil,>,'i'.iil  l.y  I^catliif^  .iiiil  hi-i  rcirtniki'iB  lu'cil  iM't.  ;is  l\;i-i 
liiciiilv  tid'ii  |iiiiii(icl  nut,  iiccissarily  imlii'iilc  Unit  mu  .li. utile  link!i);r«  liiivc  liccii  ititri' 
ilih'Cil  ill  tlic  I'iitty  jiciil  cluiiii:  it  iiriy  iiiiicl.v  in.liriiii'  tliiil  stiiHtiiiiillv  i<(iiiiiiii>  ili'ri\:r 
'iM'x  «liii'Ii  jiIkihIi  iiiiliiio  1111)11'  iciiililv  Iiavc  I'ciii  I'mini'il  Itiu'ii  i\  iliini'  nf'  i|i'.atiii:i 
liiiii  has,  Iio«f\pr,  liccn  olTcri'.l  liy  lliiitli^.v.  vU>  is.iliiltil  the  iinsatiiialcl   I'atH   aciils   (l.v 

.li»siilvin{:  tlio  Icail  siuii's  in  otiici)   fmin  |ii^;'s  tivcr  ioid  tlicii  pn" \r,\  t.i  o\iili/i'  tln'in 

Willi  alkaline  |.c>inian(;aiialc.     WIh-ii  iIh'  nl.'iii  of  tlic  di'in.f   fat   is  tlius  ticalol  at  a  1"" 
l.'iniiriatun\    twii    livilmwl    i;n.ii|is    l.i.-iinic    attai'ln'il    wln'ic    llu-   ■l-.iiKlr    linkage   cxi-tr.l 

I'dnniii^  (liii\y>li'aiii-  aii.h.  ami   ulii  r   lli.'  iiii\liirr   ii  nnw    wan I.  the  nioli'iMilc  >|ilii- 

into  two  at  tliis  plac-c,  f'.ii  iiiiii^;  two  lowio   .-oi.ls   (  ipelar;;onic  anil  azelaic)  : 

(  1  1    (11         III  Vni:<'ll(l'll    I  <'<)()ll  ; 

i  i»I<ti'   ari.i  I 

on  (III 

/  y 

,L'i   CI!       (fit  ^;     (It  I'M  "11,'  <'<MMI: 

filioNvsh^arii'  ai'iil ) 

(.■;icii,    f.'ii.i.cooii    'odii     iciiMnoii. 

I  |ii'lar^onii'   a^'i'l  i  i  a/olaii'   loiil  i 


I       .1 


KAT    Mi:TAIt«>l,l>\I 


71  :i 


Wo  imiy  nmilu'li'  fmiii  tliH  llml  tin-  ili'iilili'  ImkiiKi'  in  tlio  oloii;  lui'l  <>f  tho  depot  tat 
,aUI«  liclttvin  Ihc  iiiiilli  siii'l  t.ntli  Ci.tiHiis.  Hut  it  i«  (illi.iHiw  in  tin-  caw  nf  the  un- 
-tilumtiMl  aiiil   fri.ni  llir  IImt    i'|'i«*"">  <■'"■  "'"'''■  ''"'  "''"^■'   I'l'"'""*  "'  "li'liitiim  tliix 

vi.l.l.'il   liipiiii,'   mi. I.   wliiili,  »iii...   tills  mill    li!i«   -i\   fi 1".   wc.nl.l    in.liiiiti-  tli.if   tin' 

i|,,nl>Ii'  linkniic  rxi.lnl  L.tHnn  tli.>  hnIIi  nn.l  drMiilli  ('nlnnis.  .\n..|h,.r  jntiTrHtini;  f:i>'t 
lii.iujjlit  tip  li;;lit  l.v  til.'  i'\|,iMiini'nl;4  wns  tlint  n  ti'lr;i..\\sl.arh-  :M'i.l  nns  fnriiicl.  hIiIiOi 
I'.'ll  Hjiaii  in  «in'li  a  wiiv  a<  to  in.li.uti-  that  tlio  Ii^IiavI  y;ioii|.-.  Hniiricl  l,,.|«c.ii  llii'  sixlli 

an.l  M'vontli  an.l  l.ilu..  n  tl,c  ninth  ami  tenth  C  at -      Tl iirronw  of  this  Mnl.MtnnPf 

»oiihl  lio  s;lli»ra.-tniily   .\|.li,ilio,I    |.y  tl..'   I'ltl  o.lu.li.'U    il.lo  ill.'   I.'.'iih'  ..f  lili'ii i'l   of  II 

,„.!  .|..nl.l.'  I">n.|     i. .'..  l..lHri'n  Ih.'  ^i^lh  an.l  s.M'iiih  f  al..ni«.     •'Tho  ari.N  fonn.l  in 

the  piy's  liver  may  l.c  ac.'onnto.I    f,.i,    in   ..th.'i    «..iil«.  I.y  «u|.|.oMn:;  that    .li'wituiation 

.,r  sli'aii.-  :i.'i,|  ami  of  tl nlinaiy    (.li'|.ol  ,   ol.'ic  a.  i.l  .ci-in-'  L*    H'"  intio.lMi'lion  of  a 

.l.aiMi'  link  l.olHc.n  tho  !-ixlh  aiol  ^oM'nlh  cailKn  atonic  in  i-ach  ra-o"  (I.oathrg).  Still 
..th.'i-  .lool.h'  links  niny.  ho«.'vor.  1.,'  inli.Nln,  ..l  int..  thr  fatt*  a.l.j  .liain.  f..r  a.'i.W  of  tho 
linolic  iichl  series  mo  ioi'sonl  in  ro.l  liv.r  oil.     Kinallv.  it  is  of  inloro-t  to  n"lo  tiiat  I'npn.i. 

i„.i,l  Is  a  |.io,liirt  of  tho  al.o\o  osi.hition   | r.-.   r..i    il   has  ;'ii   .von   minil.or  of  Cat s 

an.l  thciof.MO  will  foiia  ;■(  owl.nl.Mii'  a'-hl. 

To  HO  int. I  tlif>o  .•licMiicMt  |.i'i.l)l.'iii>  ,.M.v  riii-tli.T  luTo  w oiiUl  be  out  i>t' 
jtlnci'.  One  otlifi'  raot.  >lioiil.I,  Iiowcmi-.  In-  lioiiic  in  iniiid  iinnioly.  tliiit 
tlio  tiiisattiriito.l  Ileitis  iiiiiy  lio  foniio.l  Iroiii  sutiiiatoil  Moi<ls  tlifotiRh  tlio 
iiitonncdiatc   foniiatioii  of  /i  iiyi|i'o\y   ami  /Mvctotiic   aoiils.     Tlioir  iiMfe 

piosoncc,  ill  otliiT  words,  slioiild  11..I  l.o  taken  as  ovi.lei that  tlic  oxida 

ti'.ii  of  fatty  aeids  is  iiiitiatod  li,\  tli.'  iiilrodiietinii  ..f  an  liydioxyl  -.'I'linp 
,it  tlic  (i  jjosition  ill  the  cliaiii. 


I   M 


it;;' 


i: 


I  ■  i 


n\: 


ll 


t    ■< 


lt.;i 


.  ail 


!i  i>     ' 


i 


i      i 


i'^  ■ 

-■ 

1^    * 

i' 

^ 

i^; 

(  IIAITKU   I.NNN 

<  u\ii{oi.  Id  r.Mitv  I  iimi'Kua'hui-;  am>  ii;\  ku 

TIk'  I'liiN^iiii-iitiiiii  111'  iiiiiiiiiiU  iiiiir  two  urim|r      wiiim  lil Ii'>l  hihI  i'dM 

lilniiili'il      i iinlililf  III  tliiir  ;ilillit>    In   Mlllilllilill   llii'  IiimIn    lilii|>i'iatlli  r  111 

ii  f(>n»faiit   IimI.   In  iihui'  m-  li'^^  iirliitnirv .     Ilclwi'cii   tin-  tvM'  u'riiiips  iin 
iillicr  ('xi>ts.    ii|iii'si'nlc(|    Miiiiiil.N    li\    liilifiiiiitiMu   iiiiiniiiU.    in    \v1h<'Ii   iit 

I'fi'tiiin  tiiii('«i  III'  till'  M'iir  llir  animal  is  warm  Kl |ri|  ami  at  nllici'  timi'* 

|.,,1,1M Icil      Till'  aliililv  III'  till'  liiiilM  r  mammals  in  maintain  ii  constaiil 

liiitjy  Icini'tialmi'  may  or  nnn  imt  lir  pi'isiiil  nt  tlic  limi'  of  liiilli.  Tin' 
liral-it'iriilalinir  mi'i'lianisin  nl'  tlir  Iniman  inlaiit  I'm'  i-xamiili'  remains  ill 
(l<'vi'l<i|)('(l  fur  snnir  timi'.  sn  il.al  i'S|M>snir  tu  i-nlil  is  lialili-  111  liiwi'f  tlir 
IiihI>    ti'miM'talnrr   In  a   ilallirrlnUs  (ii'iil'i'i'. 

VARIATIONS  IN  BODY  TEMPERATURE 

In  animals  in  wliirli  ll.     Iii'at  ri'milalim^'  clianism  i>  I'ully  ilcv<'|i.))('il, 

tlu'ii'  is  mil,  rM'n  iliiiinir  pi'ifi'i'l  lirallli.  i-ntifi'  ciinstaiii'y  in  ti'm;i<'ialiiri' 
in  till'  ililTi'iint  parts  oi'  ilic  1iih1>  ur  in  tin'  sanii'  part  at  ilitTi'ii-nt  iinidils 
III"  till'  iia\.  'I'lii'  aMiaui'  ni  lal  tinipiratiiii'  nl'  man  is  iisnall>  ati'il  as 
licins:  ;>"  I'.  i!iMi  !■■.  .  Inil  tin' 'liurnal  variatimi  ma\  amniiiit  In  1  *'., 
lii'liiir  lii'-'lii'sl  in  till'  lati'  al'ti'mniin  ami  |n\M's|  ilurini;  lln'  ni'.'lit.  Tlicfi- 
an-  prnlialil.v  scMial  ranscs  Inr  tliis  \ariatiiin.  ami  tlii\\  ari'  in  part  at 
least  ilepemliht  iipnii  tin'  iMeatei-  iiietalmlie  aetixitii's  nl'  the  wakiny: 
liniirs  anil  upmi  tlie  takinir  nl'  I'mnl.  Apait  I'rnm  lliese  intlllences,  linw 
I'M'i'.  ntliers  whii'h  are  less  e\ii|i'iit  appear  tn  npera'e  :  t'nr  it  lias  lieeii 
t'nnnil  tliat,  when  the  ilaily  pi-nu'iam  is  reverse.!  Iiy  niirlit  wnrk.  Ilic  tisiial 
iliurnal  \arialinn,  allhnnirh  miieh  le^s  ])rniiiiiineeil,  still  remains  evident 
e\('n  alllmnsili  this  reversal  in  lialiit  ma.\  lia\e  lieen  ke|)t  up  t'ni-  years, 
ll  is  111'  inleii'st  In  nnte  in  this  eniineetinn  that  nnetni'iial  liinls  have  their 
maximum   temperalnie  at    niiiht   ami  their  minimnm   ilnriny;  the  <la\ . 

K'ejianliii'J  the  lemperatnre  in  dilTerenl  parts  nl'  the  limly.  that  nf  the 
reclnm  is  n^nall\  almiit  1  <'.  hiirher  than  that  nf  the  nmiith.  anil  this 
afiain  hitrlier  than  that  nf  the  axilla.  Of  these  three  the  mnutli  lompera- 
tnre  is  the  must  variahle.  Inr  many  eninlitinns,  sneh  as  mmith  breatliimr. 
talkiu'.'.  (Irinkinu'  ennl  liipiiils  ami  even  exposure  tn  cold  air,  are  suffieieiil 
to    Inwer   markedly    the    tempei'atnre    of   this    reirion.      When    the    ninuth 


KiMHKI,   (IK    IHinV     II.MII  KVTI  Ml      \VI>    H  VI  l( 


715 


tcm|MTiilmv  i^  ciiiiliillv  tiiki'ii  l)y  It-ax  inif  tin-  Imlli  "■!  tlir  iliriiMmiii'ttr 
iimlfr  tlif  ti«iiuiic  \'i<v  II  iiiimitf  or  tiion",  it  is  innrtii'iilly  tin'  miiik-  ii'*  'lie 

tciii|H  rjitiir.'  n|'  till'  arl<Ti:il  M 1  "t"  tti''  liiind  wli'ii  iIm>  i-  iai>'»i'iI  '■  ili«' 

iinliiiiiiy  ••.iii.litioiis  (it'  (iiit^idc  lcm|ii'iiituif.  (inalcr  (iitTcn  tn-cs  tliiiii 
I  (',  ill  llic  tclii|MMiitllii'  III"  ilitVi'ii'iit  iiL'iiilis  "t'  the  IhhIv  nil'  ol'tcii  di 
>,i-M'.l   in    r.tl.ic  iiidividuiils  iiiid   in  lii"^'    \>illi  > iiriilat.'n    distill  1. 


FACTORS  IN  MAINTAINING  THE  BODY  TEMPERATURE 

The   l"id>    1clll|icilltuic   l.■|l|•(■^(■lll^   llic   li.l  lllinc   lictuccil    IliMl    |(roduct  loll 

Mild  liciil  loss,  Tlic  iiniiliutiiiii  is  clVccti'il  niniiilv  in  tin-  hiiim-Ics  liy  llic 
i.xidjitivc  |.r<i((ss(s  wliicli  ate  (-..Mstaiitly  ciiMiiii);  tlicrc.  Wlicii  the 
iicli\ity  (>r  the  iiiiisi-lc>  is  alwili^licd  1)\  piiiiil.v /ins.'  lli"  l.Tiuiiiiitituis  of 
tin-  iiioti.r  iicivcs  witli  cuiiirc.  tlic  IciniHiiiiniT  ot'  \  aim  liicMidi'd  atiiiiiMis 
liiiiiic(liati'ly  tails  or  liscs  accindinir  to  tin'  li'inpi'iatmi-  ot'  tlic  ciivifoil- 
iiii'iit.  A  iiliaii/cd  «  iiiiii  lilondi'd  aiiinial  is  tinis  iiiadc  In  licliavi-  like  a 
.•iildlilonilcd  till.'  Iiii'K  ascil  iiiiiM'iiliii  activity,  mi  tlic  ntlicr  liaii'1. 
lii'iiinpfly  raiM's  till'  l.odv   tciniiciatiifc  iiv   1     .  r 'J     ( '  .  iilxivc  "  liicli,  liow- 

cwr,  a    t'lirtlicr  rise  docs  not    iir.  |iidvidcd   iintliiii'j:  lias  liccii   doiio  to 

iiitcrl'cn^  "itii  tlic  iiicidiaiiisin  liy  wliidi  tlic  c\cc>s  oj"  licat  is  cot  I'id  of 
iiuiii  tlic  l)iii|\.  'I'lic  lciii|iciiitiiic  ill  Niii'li  ciM's  adjiisis  itsi'lf  at  a  liiirlici' 
level,  at  wliicli  it  leiiiaiiis  fairly  cmisl.iMt  Imwcvcr  streninnis  tlic  cxcr- 
cisc.  It  is  ))(i-.>ilile  tliat  a  certain  aiiioiiiil  i><  lieat  iiia\  also  lie  u'ciici'iitcd 
liv  the  eliciiiical  ludccsscs  occm  riii'j:  ill  llie  liver  and  other  viscein,  liiit 
ulicii  coin|iarcd  with  the  iiiiiscles  this  sniircc  nl'  licat  is  iiiidoul.tcdly  iii- 
siiriiidcaiit.  Many  of  these  cliciiiical  |iriiccssi's,  as  in  the  li\(r.  instead 
of  plddiieiiiir  actuall.v  alismh  heal,  so  that  the  halanec  lictwceii  licat- 
lirodiieiii!,'  and  hcat-cMiK  iny:  iiieehaiiisiiis  iiiav  or  may  not  chmic  ouf  in 
l'a\iir  of  the  lilicratioii  of  heat. 

The  |>rodliclion  of  heal  yocs  on  all  the  liiiie  in  tiiusi-lcs  mi  accmiiit  of 
the  ciiiidilioii  111'  tonic  emitracrimi  in  which  th"y  arc  held  see  pau'c  S14i, 
.'iiid  which  is  also  necessary  for  kec|iiiijr  the  joints  in  the  proper  doiirce 
of  llcxioii  or  extension.  When  more  licat  is  i'C(|iiircd  !>>  llic  animal  liody, 
ihc  tone  of  the  mnsidcs  increases  independently  of  the  function  which 
they  iiia.v  lie  pcrformini.'  in  emit  lolliiiu  the  position  nf  the  joints.  Tiiis 
increased  'mic  may  liecomc  so  prmiminced  thai  it  causes  visilite  cmitrac- 
lioiis.  which  we  rccoiiiii/e  as  sliiv  eriiii;.  Whenever  the  insciisilik-  liyper 
tonicity  and  the  shivcrintf  arc  iiiade<niatc  to  produce  a  siit'ticiciit  amount 
of  heat,  the  animal  instinctively  moves  ahout  in  order  that  the  •rrontcr 
contractions  may  liherate  more  heat. 

The  Ileal  is  prodiiccd  in  the  inuscics  ii.v   ii\iiialimc  ol'  llic  j'lHidsIiifis  ihaT 
haxe  liccii  assiiuilateil    fi  nm   the  Mood.      Kveii  dnriiii:  the  process  of  as- 


\   v\' 


>  M 


Ui 


y 
t 

'i 

,  1 

,1 

1 
1 

1- 

■( 

's  1 

( 

'   1 

l-ir  I 


t;:i,il|P| 


.<»:     ■ 

M  ■ 

;'j 

1* 

'!■■ 

„  ^^^' 

1^  I 

j-    ' 

li 

i  .1 


• 

1 

!■ 

'  1 

i.  , 


M 


71  r. 


Ml.lAlloI.IsM 


siiiiiliitiiiii  ilsi'ir  ii  I'iM-tiiiii  jiiiitiiiiit  (if  licMt  In  ui'iicrati'il:  tliis  is  kiioun 
,i>  tlio  >iiccitic  .l\  iiiimii'  Mftinii  nf  llio  I'iiiMlstiilV,  n\n\  is  most  i)n>n()Uii('«"l 
\\]\],  ]. lutein  :;,,il  Ira^l  ^.l  \^\\\\  imi-1mi1i\  .jiiilr  'PML'c  ."li'iSi.  Ailvillitji-jr 
iiui.N    111'   t;ik.'n   111"  tiiis  licatiii','   iiowci'  nl'   innli'iii   In   |irc>i1uce  iiiiiro   lic:it 

wli.Mi  till'  li'iu-  (Minditiiiiis  aic  r\ci'>si\  i' ;  in  winter.  I'of  cxaiiiplo,  tlu'ie 

i^  an  ini'linatiun  In  tal<t'  iiinrc  iirntciii  I'liml  tlian  dufinir  siinitiiiT.  am!  11h' 
|»'r  caiula  i'(insmii|)li"ii  <'!'  sncli  r<inii  is  iinu'li  'jii'ali'i-  in  pi'iiplcs  lixinix  in 
tfiiipi'val-'  /iini's  iVaii  in  tlicsr  Ii\  inu'  in  tl'c  trniiirv.  Tiu^  iiltiiriatc  aindiiiil 
nl"  li.'at  pniilinnl  }>y  (.xiilatiun  i^  ui-rati'sf  witli  I'af  and  least  with  rarlm 
li\  ilratc. 

Ifi„f  /r„s'  in  man  is  tiTi'. •(•'<!  paitl\  tlinmirli  llu'  luiius.  Init  mainly 
tliriinLi-li  tlic  skin.  Tlii'miuli  tlio  laltcr  patlnvny  licat  is  lost  li\-  tin'  iiliysical 
priici'ss.'s  (if  licat  (■iiiiilnclidii  ami  fadiatimi  aixl  Iiy  tlic  I'vapdratimi  df  tin' 
sweat.  Tlifdiiuli  till'  limus  it  is  lust  mainlx-  in  the  \  a]i(ii'izntion  df  tlie 
water  eiiiitaineil  in  tlie  oxpir.-d  air  Mateiit  lioal  df  vapur-.  Tlie  anidiliit 
df  Ileal  Idst  ffdiii  tlie  skill  Iiy  cdiKliietidti  and  radiation  depends  on  tin- 
teiiipeiatiire  (if  tlic  skill,  wliii'Ii  aeain  depends  mi  tlie  rato  at  \\lii('1i  tlie 
l.liidd  is  eireiilatiii'_'  tliidueli  tlie  entaiii^diis  vessels.  I'lider  drdinary  con 
ditidiis  df  e\teriial  temperature  twd  (ir  tliree  times  as  r.iueli  lieat  is  lust 
Iiy  tlicse  metliiiiK  a'-  li\  evapurat  iiiii.  Tlie  losses  Iiy  e\aii(ii  ion.  undei 
e.inditiiins  nf  rest  and  a\eraue  cNteriial  temperature,  ai-e  almllt  e(|ilall.\ 
divided   lietvveeli    tlie   lllll'js   and    tlie   skin. 

From  all  tlie^e  faet^.  it  is  evident  lliat  hat  ^-.v^■  ,h-ri(i:i  iiniiiilii  lui  II" 
J,  ,;i  ,/()(/  fulfil  III  II  utiiiill  ilii/rir  hii  llir  liiinin.  Tliis  iiKMiis  tliat  under 
a\('ra!.;c  ediiditinns  in  man  //"  uniiii  n  i/iihilimi  uf  halt  lu.^^  Is  rljcrlnl  hii 
riir!iiliii!is!  ill  llii  sl.iii  h  ni ju  rul uri  Iii  nitrlit  alinut  Iiy  periiilieral  vaso-eoii 
-.trietimi  and  dilat.ilidii.  'I'lie  iiiaiked  sensitivity  uf  tin'  eiitaiicdils 
lilddd  supply  tn  .-liaii'ies  ill  the  temperature  of  the  environment  has  heen 
\erv  eleaih  sliuwii  li\  ■iliser\  a'iuiis  luade  with  the  hand  ealdriiiieter  uf 
Slewarl    deseiilied    elseuhele       pa'je    L'^  T  .       When    til"    lild.idtlnu     tlirdll'jh 

the  hand  is  eNaniineil   in   a   peisun   whn  has  lieen  exposed  to  the  diltside 

,iir,   it    may  lie  little  i c  than   half  that    whieli    it   attains  after  he  has 

lieeii  ill  a  warm  iddin  fur  suiiie  time  In  the  diitside  air  the  vessels  cdii 
strict  to  prevent  lii-at  loss  Ii\  ■■i.nduetidii  and  radiation:  in  the  warm  rdom 
thev   dilate  td   facilitate  this  l,,ss.     The  afferent    imimlses  wliiidi   rellexly 

.•niitrdl  the  chaii'^e  in   the  cntaiiediis  M 1  cirenlatiiin  may  he  set   n]i  I'.v 

Ideal  applii-atiiiiis  ,,f  heat   nr  i-i^^i\.  as  can  lie  sh.iw  n   in  the  Iiaiid-ealorim 
eter  eNperiiiieiits  liy  aiiplviii'/  a  culd  iiad  tn  the  skin  df  the  edrresixiiid 
\\\<S  fdvearm,  when   an    immediate  cnrtailment    df  Iihiddflovv   takes   pltice. 
<ir  the  vedex  may  lie  excited    from   distant    skin    areas,  jis   illustrated    in 

tl iirtailliient    ill   Moodlhiw    dliserved    wlieii    tlie    npiiosiie   hand    to    lli.il 

dii    which   the  dli-(M-v  ;it  imi    is  lieinu'   made   is   placed    in   cold    water.      Th' 


(ciN'rum,  i<y  ikihn     i  i  \init  \i  i  isi     wn  iiam; 


iiih  i>  niNi'ltli 


717 


•pcllil 


Ilia^llilUili'  <ir  llli'  "•liillli'i'  ill  i-nIilllriMI>  .■Il.-Ul;illnh  1>  llrVlTIlH-lc; 
,.|i1  Ulioli  llif  fXt.Mit  oi  the  inv;i  of  til.'  l.n.ly  tliiit  is  ..|.|)OSimI  to  tlir  .■lillli'.'c 
ill  tcinpfiiiHirc,  as  sen  in  tlu'  iliialiitinii  of  tin'  skin  vcss.'ls  ptioi'  to  a 
rise  in  lioily  Ifnipi'i'iitiirf  wlicn  a  |i<Tson  is  imniciscil  in  a  waiiii  Iialli. 
AltlioiiMli   atV.'irni    iiiiiniiM's   iicni   tlir  -.kin   .n-v  tiu'ivl'oiv  oi'  ■rrent   im 

IHn-laiicr    in    a.i.iiisl  iiiii'    tlir    .■utaiifmi.    1,1 1    >n|.ipl.\     a r.iin-;'    to    tli^ 

aiuount  ol'  slift'ai-f  i iiinr  that  lias  tn  n,-,-iir.  ,i   I'ui'tliri'  fircTt  is  also  pio- 

(luccil  on  tlK'in  liy  tlic  artioii  on  iIh'  ii.i\o  criitrrs  ol'  tciti|)cratiirc  dif 
I'crcnfcs  in  tlic  Mood  itself.  Tims.  \\  li.'ii  ilir  tcnii'ffatniv  of  hlood  uroinu 
to  tlif  lifaiii  is  raiM.l  l>y  placinir  tin'  caicli'l  artfiics  on  some  lieatin^  <le- 
vieo  Of  wlieii  tlie   l-e-ion   ..f  111.'  .-.irp.na   striata    i>  .liveetly   wanne.l.  111.' 

skin  vessels  li.'eoni.'  .lilate.l  as  if  tlie  aiiiiiiiil   had  I n  .'Xi.osed  l.i  'j:''1i<'I-!i1 

w  aniilli. 

When  ill.'  l.iss  .if  li.'at    !■>    ra.liat  i.ni  nii.l  .•..iidii.'tioii   is  no  loipz.'!'  ad.' 
.piate  t..  iii'event  a   rise  in  l".dy  t.nip.-ratiire.  ..i-  wli.'ii   tli.'  j.r.ie.'sses  fan 
n..t   op.'iate  on  a.-. -..tint   ..f  a   lii'Ji   t.'iiip.'i-atu'.'  in   tin'  .'iivii-onni.'nt,  the 
l,,ss  .if  h.'at    from  the  skin   is  mainly  .l.'p.'ii.l.'iit    upon  //..    ,  rapnnilion  of 

•iinal.      rnd.T  onliiiary   litioiis   this  .'vaporati.m    tak.'s    pla.'e   at   su.'h 

a  rat.'  that   th.'i'e   is  no  visihl.'  p.'i'spiralion  on   the  surfa.'e  .if  th.'  liody 
Ih,'  s,.-.-all.'.l  inseiisihl.'  p.Tspiral  i.ui.     Winn  th,'  lii'at   loss  l.y  this  ehanii.'l 
must    he.'ome  jrr.'ater.   tli.'   in'i'spiration    is   pro.liie.'.l    in   larjr.'r  aiiDiliit,   si. 
that  it  .'olh'.'ts  .111  the  snrt'a.-.'  ..f  th.'  ho.ly  :  an.i.  i>rovide.l  the  .•on.litiotis  ..t 
the  .'iiviroiimi'iit    ar.'  such   that    evaporation   .-an    r.'a.lily   take   plaee    <h>\\ 

relativ.'  hnmi.lity   ,  th.'  aiiionnt  ..f lini;-  <•(  th.'  h.i.ly  that  .'an  h.'  elTeet.'.l 

hee.mii's  very  !.'reat.  A  man  nia\  exist  with. nit  any  mark.'d  lis.'  In  body 
t.'iiiperatiir.'  in  a  very  hot  enviniiim.'iit  .'Veii  wh.'ii  h,'  is  exposed  t.,  an  out- 
si. le  t.'iiip.'ratur.'  tiiat  is  the  saiii.'  as  that  ..f  his  ho.ly.  .n-  .'v.'ii  .jr.'at.'r.  T., 
.■in-:'iirau.'  I'vap.iralioli.  h.iu.'V.'r.  h.'  sh.'illd  !..■  naked  >>v  \.  r,\  li'uhtly  .'lad. 
anil  the  air  shoul.l  he  ki'i>t  in  •■oiistant  moti.ni  so  that  the  layers  of  air 
next  to  th.'  skill,  whi.di  ..r.linarily  vry  .|ui.ddy  li.'.'om.'  satnrale.l  with 
vap.U',  tire  traiisferre.l  aii.l  r.'pla.-e.l  h.x  .lr>ei-  air.  .Mov.iii.'iit  of  th.'  air 
als.,  iiier.'ases  the  heat  l.iss  l.\  eniidii.-tion.  pinvided  th.'  temjierat  ure  of 
tlie  air  is  not  too  near  that  of  the  hody. 

The  importaneo  of  th.'  ni.iv.'inent   >•(  air  in  the  ri".;iilatiim  of  heat    lnss 

hasli.'.'ii  el.'arly  .l.'iiionslrated  li.x    I nard  Hill.  '  K.  S.  I,,  e.  and  ot  li.'rs.  n\  h- 

have  fotin.l  that  a  L'r.'at  ].art  of  tli.'  ilis.',.nifort  .'xp.'ri.'iiee.l  l,y  liviu-j:  in 
stagnant  air  .'an  li.'  ohviate.l  h.v  puttiiiir  tin-  air  in  m.iti.'n  hy  .'leetrio  fans 
with.iut  .loiin^  anythinir  to  im|,i'..v.'  its  eheiui.-al  pui'ity.  In  one  famous 
oxperiineiit  a  innnher  of  yoiiiie'  iii.'ii  wer.'  pla.'.'.j  in  an  air-ti'.dit  cabinet 
at  the  ordiiniry  l.'iiiperatur.'  ..f  th.'  room.  Afl.u'  a  time  they  began  t.. 
exhibit  Ih.e  svi!!(>t.i!!is  usiudly  attrihute.l  i.i  pollut.'.l  air:  they  beennn' 
.Irowsv   an. I   s.uii.'   <>\'  them   .l.'veloit.'.l   hea.laehes,   .'t.'.      A    small    cleetrie 


?  ■ 


ft' 


w 


u  w 


m 


71S 


Ml    r  Mini. ISM 


ii  nihil 


:i    Hi    : 


fl». 


1  1 1 


■  i;  ■' 


4if^h 

1    ;. 


r-^  i 


fiin  wiis  tlit'ii  Nliii'lnl  Ml  ;is  to  si'l  the  .•lie  in  iniilidii  iiniiu'diiitt'ly  all  m 
the  iiKMi  rocovcit'd  iind  ri'iiiiiiiiccl  in  :i  pcrt't'cllv  ciiiiit'orlaMc  conditioii 
so  loni;  as  tlic  1'aii  was  ki'i>l  (idiinr.  'I'lic  itractical  aiiplicafii'ii  of  llu'sr 
t'ads  to  fill'  livfjii'iiir  I'oiifrcil  (if  llic  w  cirUinir  I'Diidilinns  in  ininc  sliai't--. 
ill  snltinai  incs,  in  workslmps.  itc.  will  lir  self  i'\  idi'iit. 

Tilt'  Ntiiiiiiliis  1(1  iiici'ciisi(|  sucatiiiij  scciiis  t(i  lie  (jcpciidciil  mainly  mi 
cliaiiiris  in  till'  t(  iiipcialiiic  nl'  tlic  lilodd  :  I'ln-  sweating  dues  not  iiii 
mediately  set  in  when  tlie  liddy  is  siilijected  t<i  lieat.  as  liy  a  warm  liatli  nr  a 
lidt  jiai-k.  It  usually  takes  I'ldin  ten  In  t\v('nty  minutes  after  tlio  pevsnii 
has  been  jilaeed  in  the  liatli  ni'  siiridiliKled  liy  the  warm  lilankefs  of  tlic 
pack  liet'ore  sweating;  Iieeomes  prdiidiineed.     It  eaii  usually  lie  slidwii  that 

liefdie  it    sets  in   the  luidx    l(  iiipciatiiic  lias  1 n   raised    from  O.l  to  ()> 

dejrrees  ( '.  i  Oil  to  1.4  de^rrecs  !•".  .  In  this  rcjrard,  tlierefdic.  the  response 
df  the  sweat  ;daii(ls  ddcs  not  dcciir  so  |ii(implly  as  dues  the  dilatation  of 
the  eutanediis  xcssels. 

I,oss  df  heat  liy  e\  apiiiat imi  df  sweat  iirs  only  in  certain  animals 

It  is  ]ira<'tieally  aliseiit.  mr  e\am|ile,  in  tli(>  do^.  The  de-rree  to  Mliiidi 
it  may  dceiir  also  varies  in  dilTereiit  individuals  of  the  same  species.  The 
]i(iuer  (if  w  ithstaiidiiii;  hiuh  temperatures  is  prupditidiial  in  man  td  the 
facility  with  whi(di  he  piispires.  Where  sweatiiii;  is  interfered  M'itli  li\ 
skin  diseases,  liy  iclitlivdsis,  fdr  example,  exposure  to  lioat  or  in 
creased  heat  produelidii,  as  liy  muscular  activity,  may  raise  the  lind; 
temperature  td  a  daimcrdlls  dej;rce. 

Alidther  factor  upiin  which  the  cflicieiicv  of  e\apdrati(i|i  (if  sweat  in 
cddline  the  lidd\  depends  is  the  relali\c  humidity  of  the  air.  Wlieii  this 
is  hiiih.  c\  apiiratidii  df  water  intd  d  can  not  occur,  and  it  is  on  tliis 
accdunt  that  an  increase  in  Imdy  teiiiiierature  is  mindi  more  likely  to 
dcciir  in  warm,  humid  atmiisplieres  than  in  those  that  are  <lrv.  At  the 
same  tcinpei  ature  iieojile  can  live  in  perfect  cdiiifort  in  the  dry  air  of  the 
iipeii  )ilaiiis,  lull  siilTcr  immediately  frdiii  rise  df  lemperatiire  when  they 
ud  into  the  humid  air  nf  the  rixcr  \alleys.  Similarl,v,  work  in  hot  fac- 
tdiies  (ir  ill  mines  is  (piite  piissilile  at  very  liiiili  temperatures  if  the  air 
is  kept  dry  and  in  motidii.  hut  liecdiiies  impiissilile  when  the  air  is  moist 
111  judiriiivr  of  the  ade(|uacy  of  air  from  this  point  of  view,  it  is  tliere 
fore  iniport.iiil  to  take  not  the  (inliiiary  dry  liuHi  tlierniometei'  readiiii; 
lull    that   df  the  wel-liillli." 

Ill  animals,  like  the  den;,  that  dd  iKit  jicrspire  over  llip  surface  of  the 
liody,  rnixirrjiliitii  nf  llu  irah  r  in  lln  i.iitinil  iiir  is  the  most  important 
iiiethdd  df  rciiiilatidii  df  heat  loss.     When  siudi  an  animal   is  exposed  t" 

■Ihr     ...lliili,    111,  in,,..  1,  1.  r     1,^1-1.1-     ,1     1,  ini...i:|liii.-     (li,!     I-     !•.«..■     lli:...     ill, I     ■■!     Ilio     .iry-ljulb     I!, 

I  1  ,il„>rtii,M  ti,  ill,-  itl.iliM  !i,.ir,n,!nv  ,,t  llu  .11:.  W  lu  11  llic  ;iii  1-  niiTiii]c!c!\  s.ttiiiiitiil  with  innisturc. 
tin-  tfni(ii'r;itmr  r,i,,r,U,l  l,v  tl,,-  t.x,,  niviimiH  rits  will  1„  tlir  v.un,  ;  \\!,ir.  it  i-  inrf'Tllv  ilry,  tlu 
liiffrrencp  will   lie  ;-.:iviinal. 


(UNTKUl,    III'    linllV     ir.Mn  HATl  Hi;     \\l>    IIMI! 


719 


uiiiinth  or  wlifii  tlic  irtiioii  of  llif  cdrptira  striata  is  artiticialiy  wanin'il, 
llii«  broatliiiiK  ininiodiatcly  lu'comes  much  (|ui('kor  and  dcciu'i',  so  that 
puliiioiiip  ventilation  is  <;i(>atly  iiicroasod  and  much  more  water  is  carriod 
out  ns  vnpor  with  the  expired  air.  To  vajHirize  the  water  laiRe  quanti- 
ties of  heat  are  retpiired  (seen  in  the  latent  heat  of  steanil.     Tn  man  this 

■,.  .,  .        ..  ,     ■ 1  ...     :i     l.„..^...,„    ^.. 


I'thod 


method  is.  ordinarily,  not  of  <>reat  iinixntiini'e,  hut  it  may  lieeome  so 
when  sweatinir  is  interfered  with,  as  in  iehthyosis.  The  more  rapid 
lireathini:  also  faeilitati's  eoolinir  hy  incicisini.'  tne  eoixhletion  of  heat 
t'ldiii  the  niueous  iiiemtiraiies  of  tlif  tonirue,  month,  throat,  etc.  The  im- 
poi'tanee  of  this  mi'thod  of  cooliii'^  has  lieeii  shown  liy  findinjr  that  after 
the  iutroduetion  of  a  liaeheal  eaiinula  a  <loi;  ean  not  withstand  an  in- 
crease of  external  temperature  nearly  so  wi'll  as  a  normal  animal. 

There  are  man.v  other  (pii'stioiis  eoiicerniin.'  thi'  i-oiitrol  of  heat  loss 
from  tlie  liuman  iiody  that  iniiiht  he  considered,  hut  it  is  scarcely  ne<-- 
essary  to  do  so  he'c  It  shouhl  he  added,  however,  that  the  relative 
humidity  of  the  air  in  the  control  of  heat  loss  has  a  different  sijrnificaiirc 
when  the  temperature  is  hijih  from  that  when  il  is  low.  Tli-ih  relative 
Immidity  at  hii;h  temperatures,  as  we  have  seen,  interferes  with  evapora- 
tii.ii  of  sweat,  whereas  hi-h  relative  humidity  at  low  teiii])eratures  in- 
ci'eases  the  hcat-condnctiii!:  jiower  of  the  air  and  th.'rcfore  tends  to  cool 
ofl'  the  surface  of  the  hody  by  irreater  condnclioii.  11  is  on  this  account 
that  it  is  much  nnn'i'  comfortahle  to  live  at  a  low  tempeiatiue  when  the 
air  is  dry  than  uheii  it  is  moist.  <»ii  the  <lry  jilaiiis  of  the  West  a  tem- 
jieratiire  of  man\  ileirrees  helow  zero  causes  less  sense  of  cold  to  he  ex- 
perienced than  in  the  moist  atmos|.liere  at  a  coiisiilerahly  hijiher  tem- 
perature alonji  the  Crcat   Lakes  and  in  the  river  valleys. 


THE  CONTROL  OF  TEMPERATURE 


,oln 


In  the  ease  of  man  the  hody  temperalme  is  \eiy  largely  nndi'r  vi 

tary  control,  as  hy  the  clioi if  clothinu'  and  the  artilicial  heating  of  the 

room.  Desirahle  as  this  voluntary  control  of  heat  loss  may  he,  tliere  ean 
he  little  dimht  that  it  is  often  manajred  to  the  detriment  of  liood  health. 
I.iviiifr  in  overheated  rooms  dnrinu'  the  cooler  months  of  the  year  so 
diminishes  the  loss  of  heat  from  the  hody  that  the  tone  and  heat-)i!  odnc- 
ins,'  powers  of  the  muscular  system  are  lowered.  .Vot  only  does  this 
diminish  the  lesistance  to  cold,  hut  it  causi's  the  food  to  he  incompletely 
metaholize<l  so  that  il  is  stored  away  as  fat.  The  superfieial  capillaries 
also  heeonie  eonstricted  and  the  skin  hloodless  and  "pasty."  It  is  not 
looks  alone  that  sufi'er,  however,  hut  health  as  well,  for  hy  havinsr  so 
little  to  do  the  heat-rerrulatinu:  ine>-hanism  uets.  as  it  were,  out  of  gear. 


'     1 


tit 


M 

I  n 


!     f 


■Ui^'    I 


It' 


i 

!1 

1   ■ 
i 

i 

if' 

tl 


I  , 


1 1 


1 
' 

t 

m 

liS. 

^ 

7-J() 


Ml  IMiiiMsM 


M.  Ihal  uln'ii  it  i^  rc(iuii-.'<l  In  .-ict,  ;is  wlicii  llic  |p.'lsiin  jicics  dutsidi'  I" 
tin-  t'oltl  ail',  it  iiiiiv  iiiit  i|i>  M'  ;i>  iiioiii|ill>-  ji-i  it  sliotild,  witli  tlie  rt'sult 
that  tlic  l)t)(ly  ti'iii|>t'iatuic  falls  Minicwliat  and  cafari'li,  etc.,  ai'c  tin 
ivstilt.     Tliiif  can  111'  little  limilit  tliat  iniicli  ol"  the  boncfit   of  opeii-aii 

s!('i'|iiiijr   is  ciwiiiiT  tn  till islaiil    stiiiiiilalinii   (if   ilii"   iiK'taliolii'    lU'occssrv 

M  liii'li  it  I'ausi's. 

As  will  lie  iiifi'iicd  fiiiin  w  lial  lias  liccii  said  almvt'.  llic  (Mintinl  between 
lieat  prodnetiim  and  lieat  lnss  is  etVccted  tlii'oiiyli  a  ik  m  vinlir  lorated 
in  di'  near  111/  eiir|>iira  striata.  In  nmsl  animals,  when  the  siiinal  eoi'd 
is  cut  in  the  eci\ic:il  fcuiim.  tiic  lindy  tcnii)efatiife  (piiekly  falls  nnlcss 
aitilically  maintaincil.  In  the  case  nf  man.  mi  the  othef  liand,  it  l\a- 
usually  heeii  dliscixi'd,  after  ai'cidental  sei'timi  of  the  spinal  eufd  ill  tin' 
cervical  reirinii,  that  a  lise  in  temperature  nccurs.  In  1\\cnty-foiir  uii 
eiiiaplii-alcd  easels  of  spinal  curd  injury  in  man,  enllected  frnni  llie  rec 
ords  (if  (luy's  Ilii>>pital  by  (Jardiner  and  rembrey,  it  was  fmind  Dial 
nineteen  showed  hypertliermia  i  sonieliiiies  amounting  1o  4:5. !>  (".),  and 
only  live,  hypothermia  i  sometimes  'JT.ii  ('.  .  Tf  the  patient  lived,  the 
ultimate  elTeet  of  the  section,  as  in  the  lower  animals,  would  no  doub^ 
be  the  loss  of  the  jiouer  of  maiiitainin'.r  a  constant  temperature. 

The  extei-.t  to  w  Inch  the  animal  co s  to  luliavc  as  if  cold-blooded  aftei- 

section  of  the  sjiinal  eoid  varies  considerably  according'  to  the  level  oi 
the  lesion;  if  the  cord  is  cut  in  the  u|)|)er  thoracic  re<;ion,  for  example, 
the  rciiulation  auain^t  cold,  allhouirli  distincll.x  less  eflicient  than  normal, 
is  far  lietler  than  ^^hen  the  section  is  made  Ihrotiirh  the  cervical  cord 
This  difTereiice  is  de]icndcnt  on  the  fact  that  after  the  lower  lesion  much 
tarL'er  muscular  liioups  and  skin  areas  are  left  intact,  so  as  to  make 
ri  Siulation  |ios>ible.  Section  of  the  dorsid  <-ord  in  mice  has  been  found 
by  rciulire>  to  abolish  entirely  the  increased  metabolism  which  neciii- 
in  normal  mii'e  wIkmi  they  are  e\)ioseil  to  cohl. 

In  the  lifjht  of  these  expeiinieiits  it  is  luoliable  that  the  dilTerelice  in 
ill  .  ,  Is  ))rodni-ed  on  body  temperature  bv  section  of  the  eervica! 
spi,.  .rd  in  man  and  the  lower  animals  <lepends  on  the  relative  iiii 

|)ortance  of  the  licat-i)rodu(  iiiii  and  heat-dissipatinj;-  mechanisms.  When 
the  control  c.f  heat  loss  is  jiaraly/ccl  in  the  smaller  animals,  the  eooliii'i 
of  the  bodv  becfimes  excessive  in  relation  to  the  amount  of  Inat  produced 
in  the  jiaralyzed  ninscles,  because  the  body  surface  is  extensive  in  com 
parison  with  the  bod>  weijrhl  i  see  iia^re  .'i.Mi.  In  the  lai'srer  animals  sudi 
as  man,  on  the  other  hand,  the  coolin>r  effect  is  much  less  marked,  esjic 
cially  ^^■hen.  as  is  comnioii  alter  such  an  accidinit,  the  jiatieiit  is  kepi 
unusually  warm. 


iipNTItol.   UK    llcilvV    'lIMlTinTrm      \M>    KKVIR 

FEVER 


721 


III 


The  clinical  (ipplicati.m  ..f  a  kiiowlcliic  of  the  incchnnism  of  heat  rcsrn- 
latioii  in  the  animal  ImmIv  coiicitiis  llic  causes  of  fever.  In  the  most 
familiar  form  fever  is  pro.liici'il  l.y  infectious  i)rocesses,  l.ut  it  may  also 
be  owiiisr  to  various  other  causes,  ainonir  wliii-li  may  he  mentioned  the 
parenteral  injeclhui  of  foreiirii  protein,  excessive  .lestructiou  of  protein 
suhstances  in  the  ho.ly  itself,  the  iictiou  of  certain  drUKs,  an.l  lastly, 
injury  to  the  base  of  the  hrain  or  lesions  of  the  upper  levels  of  the  spinal 
cord.  Various  types  of  fever  are  rccoijiiized:  when  the  temperaHire  re- 
...aiiis  constantly  above  the  normal,  it  is  known  as  continuous  fever: 
uhen  oscillations  oceiir  but  the  tcmi.cratuic  never  falls  to  the  normal 
level,  it  is  known  as  ri'iiiitteiit  :  wlien  it  .Mttains  the  normal  level  at  cer- 
tain periods  duriiiu'  the  day,  it   is  known  as  intermittent. 

Causes  of  Fever 

Durinj;  a  stiddai  rist  in  tem|ieiatiirc  there  is.  on  the  one  liaiiil.  in- 
•■roased  heat  production  in  the  muscles,  an.l  on  the  ether,  diiuin- 
ished  heat  loss  from  the  surface  of  tbc  iKxly.  rh<  (<  r,  r  Ix  thnuforr 
ihi(  to  (1)1  rxainiirnliiiii  of  tin  /)(•(/<■< s.-.v >■  l>ii  wliirh  tin  lioihi  mn-malht  rc- 
mts  to  roiKUtioiis  which  tf)i<l  to  loir>  r  thr  1io,hi  t<  nipi  rnliirr.  The  increased 
muscular  activity  thus  induced  often  causes  visible  contractions,  familiar 
as  shiveriiii::  ami  the  .•onstrictioii  of  the  cutaneous  blood  vessels  pro- 
ihiccs  the  sub.jective  sensation  of  chills,  and  causes  ihe  skin  to  become 
pale  and  cold  to  the  touch.  The  skin  muscles  also  contract,  producing; 
••i,'oose  skin.'"  Duriiiu:  this  stairc,  ob.jective  denioiistration  of  the  cur- 
tailment of  the  skin  circulation  can  Ijc  sceure(l  by  observation  of  the 
l,|no<lllow  tiii'ou-h  the  hi.nils  an.l  feet  '  pa-,-  -js:!!.  When  the  ti'iiiperatuiv 
M«l<lnil!i  lulls  .if::\\u.  the  crisis,  as  it  is  called,  muscles  b.'come  llacehl 
and  produce  less  heat,  and  the  cntaiiciuis  blood  vessels  dilate,  as  has 
lieen  shown  by  measureiucnts  of  the  blo.idllow  of  the  hands  and  feet. 
M  the  same  time  also,  the  swent  inlands  are  stimulated  and  marked  per- 
spiration occurs. 

Coueerniii^  //"   "'".v    "/  coiiliiiiions  fi  r,r.  it   must   b.'  assumed  that  the 
lialauee   between  h.-at    i>roduclio!i   and   lic.-it    loss   Iwis   been   ad.just  t    a 

hiuher  plane  than  normal.  \Vc  can  imt  ,-xi.l;iin  th.'  fever  on  t,  asis 
citlicr  that  heat  production  is  pci  in.nicnll\  incicised  '.v  lliat  heat  loss 
is  iiermanently  diminish.'d.  for  in  ncithci'  of  tlies,.  ,-as.'s  would  the  tem- 
perature stand  at  a  i.crmancnt  level  but  v.ould  steadily  rise  or  fall,  ae- 
cordinsr  to  which  mcchanisiu  was  ilisturbed.  While  set  at  this  hisrhcr 
plane  of  fever,  the  thermosicnic  ioiv.  .ciiicrs  ;irc  stiU  cap-nble  of  ve 
spondins;  in  the  usual  way  to  the  influences  which  cause  the  body  tern- 


:  I' 


m 

•  1 

i'  ■;! 

^1 1' 

til. 


U'l 


1 '  iir 


!::;■ 


ill 


!  ^' 


'1:1  l! 

,     i 


iH«^ 


7-'i' 


Ml. I'M  ml. ISM 


piiiitiii r  tci  cli.-iiii;!'  ill  ii  iioiiiimI  iitrsdii.  Vor  cxaniplc.  wlicii  a  iVvcr  i>m 
ticiil  i>  >iil).ii'cli'.|  tci  a  Imt  liatli  m(  tliat  liis  Imdy  1cin]i(>rafur('  risos  iiliimi 
0.2  to  0.."t  cli'jiii'c's  ( '..  swraliiiji  (ici'iirs  .just  as  in  a  iKirinal  indiv  iiliial :  nr 
if  cxiMcivi'  is  tiiki'ii  tlic  iiirrcaMMi  aiiiinint  of  licat  tlicrcliy  iirodiiccil  in 
till'  iiiiisfli's  is  (lissi|)atiii  ill  till'  Usual  wav.  M'licii.  mi  tiio  ntliiT  liaiiil, 
till'  patii'iit  is  I'xpiisi'il  til  riilij,  the  \  isscls  iil'  tin'  skin  niiitrart  ainl  Ih' 
sliivi'is. 

Altliiiiiiili   lV\i'i-  is  imt  raiisi'il  liv  an  ai-tiial  ilistiiiliain-o  nf  'lalani'i'  In' 
twi'i'ii   liiat    luiiiliictiiin   aiii!   lirat    loss,   ncitlicr  of  tlicsc  iiroi'rssos   is  l>iii 
i-i'i'ijiim-  at  its  iKniiial  1i'\rl.     That  tln'ir  is  a  ilistiiirt  iin-ri'asi'  in  tlio  tntal 
lii'at    iMiiilurtidii    111'   ilif  liiidy   in   ai'iiti'    iVvcis   in   \\  cll-ilfvi'ldpcil    persons 
lias  lii'cii  sliiiwn  liy  im-aiis  nt'  tin'  ii'spiiatimi  caliMiiiirtcr.     Tliis  iiifToasi'il 
lirat  piiiiliii'tiiiii  is  iii't  iilisiTv  I'll  in  |)atii'iits  hIio  lia\('  lii'i'ii  linmalit  iiitu 
a    w  cakcinil   (•(niilitiini   and    in    «lioiii    tin'   iiiiisiMilar  tissiii's   liavi'   IiiM-nnn' 
at  lopliii'd   Ky   liiiii;  I'liiitiiiui'il    Irvrr.     Tlic   inci-i'asrd    licat    prndiii'tiim    in 
I'lintiinmns    I'i'Xit    is   niainl.v   di'pi'iidriit    iipiui    tlii>   iiicn-asi-   in    Imdy    ti'iii 
pt'raluir  and  is  nnf  niir  i>\'  its  i-aiisrs,  ;is  is  i'\  idcnt  I'nun  tli(>  fact  tliat  far 
larui'r  i|iiaiil  llii's  nl'  lirat   ari'   l'ii'ipiriitl\    piudufi'd   in   iiiiriiial   individuals 
as  a    I'l'siilt    "I'   iiiiisi'iilai-  cMTrisi'   or   tin'   takinir   "f  laru'"   (piantitii's   i<\' 
pidti'in  riidi    IhimI.      Till'    lii'at    lliiis    pi-nduii'd    is,    Imw  i'\  it.    vci-.v'    quicki\ 
dissipati'd,    SI)    tliat    uiily    a    trnipiiiar.N'    ii>i'    in    ti'iiipiTatufi'   iin-urs.    (r{. 
Hi'wli'tt.'"^ 

Siiiiilail.v.  it  ran  In'  show  n  that  in  ciiiitiiiuiius  IVvim'  tlicri'  ;..  a  rclativi' 
ini'llii-iriii'.x  in  the  iin'rlianisiii  of  hiat  dissipation.  Wlioii  tin'  ti'inporaliiri' 
of  a  iiiiiiiial  piTsiiii  is  aitiliriall;  raisid  thioui:li  almut  1  ('.,  a  inafkcl 
iniTrasi'  in  i-iitaiiroiis  liloodlli;\x  and  profiisi'  pcrspiiatinn  an-  invai'ialil\ 
iiiitnl.  In  a  patiriit  with  fi'v  it  of  thr  saiiii'  di'jiriT.  on  tlic  oIIht  liaiid. 
tlit'i'i'  is  piai-tii-all.".'  no  rhainii'  in  tlic  skin  ciiciilalioii:  indeed,  it  is  usiiall.v 
diiiiinisht'd,  and  there  is  no  unusual  piTspiration.  The  lii'at-rey:iilatiirj 
iin'chanisiii  is  t\u\\  liNcd  on  a  plane  that  is  hiirher  than  the  noi'inal,  so 
th:  '  alllinuy:h  fuitluT  iiicreasi'  in  lioily  tciiipcrature.  as  we  ha\e  seen, 
calls  ;.,ith  responses  like  those  in  a  iioiniiil  iiidi\idual.  .\  et  at  the  fe\  er 
ti'iiipei'ature  itsrif  there  ;ire  none  of  the  reaetiiins  which  a  iioniial  iiidivid 
iial  would  cxliihit  if  his  leiiipeial  lire  were  artiticiall.v   raised  In  that  level.  ' 

The  ad.jllst  nielit   of  the  temperature  at   the  liieher  level   is  hy  no  means 
Ml  perfect  as  it   is  at  the  normal  level  of  health,  so  that  a  iinrmal  suli.jeet 

is  mole  resistant  to  tl iTcets  of  I'old  than  is  a  iiatient   with  fever.     The 

deerec    of    response    of    the    fever    patient.    llo\M'\ el',    \aries    consideralil> 

fioiti  time  to  time;  : Id  hath  in  ty])lioid  fever,  for  examiile,  lowers  the 

!iiid.\  teii;pei,ittiii-  iriiiel;  li-ss  effeetively  at  an  early  s1a<ie  in  the  disease, 
when  the  fi'vci-  is  more  or  less  eontiiiiinus,  than  later  w  lien  it  is  heeoniiii"^ 
of  the  intcrmilteiit  t\pe      In  the  third  week  of  the  disease  the  cold  linth 


iNIKill.    (l|-    I'.dhV     11   MI'lltMl   lil       \M>    I  I  Ml! 


T>-\ 


iiK.ii'  iciulil.N  liriiius  <l«iuii  \\\r  tciiipiTMtiiii'  iiiiil  kci'iis  il  ilowii  I'lif  ii  lipiiL'ir 
liisio  limit  iluriii«  the  fust  in-  mmmuhI  w.M-k.  The  iiit'cli.uiisiii  fur  licat  lnss 
is  Ills..  (Icrniincil  in  I'fvcr,  wliidi  cxiiliiiiis  llio  iis.>  in  ifniii.TMlinv  lliat  is 
likclv  to  fc.lldw  till'  iM'irnriniuifi'  .if  .'Vfii  iii.Micr;it.'  nni^.-nliif  I'x.'i-.'isc  ..r 
llic  liiUiii'.'  i.f  tiM,  lii'iivtv  il  nH;il  ill  tiiliiT.'iilnns  mikI  .'imv  aL-si-.^nl  tyiili'mi 
liiiticnts. 

Changes  in  the  Body  During  Fever 

111   scckiiiL'^    fnr   th.-   cnils.'   <.f   f.'\fr   wlii.'li    is   (\i.|.ntiy    <.f   an    ulis.'Ui-.' 
nature,   it    !■>   niM-csNiu-y   t.>  .-.ilj.'cl    all    ill.'   infuniiatinn    u  .•   '-an    n"_'ar<liii!r 

til.'  in.'taiii.lii-  cliaiiKi's  tliat    ar.'   tlifi cuiriiiir   in    tl;.'   animal   Imdy.      A 

few  ..f  111.'  must  siniiilii-aiit  fa.-ts  tliat  liav.'  s,,  far  Im.'II  .•..llfctcil  may 
lie  iii.'nti..ni'<l  Ii.tiv  Si. mi.'  «(  tin-  iii..st  iiiiii..itant  .-..n.-.'ni  tlio  .lis- 
Iin'liaii.-.'  ill  iiiti-..!icn..Us  ciinililn  iiiiii  .•alls.. I  l.\  lli.'  ...nsi.l.'ial.lf  loss  ot 
nitr.i._M'ii  ulii.-li  tak.'s  |ila.-.'  in  f.'v.r  |iati.'ii1s  v  l,.'ii  ili.'y  ar.'  iV.l  on 
tlie  Usual  li.ispital  (lift  )ir.'s<-iili.'.i  t'..r  sudi  .'as.s.  This  J.iss  ..I"  iiitro- 
u'.'ii  is  no  <l..nlit  tin-  r.'sult  of  the  partial  starvation  in  which  tli.'  pa- 
ti.'lil  is  k.'l>t  ;  for  il  has  Im-.'H  slioun  l.y  Shal'f.'r  and  Col.'iiian  that 
pati.-nts  with  lyplmi.!  \'v\r\-  iiiav  I..'  maintain. •(!  in  nit  r..<j.'ii..us  ..ini- 
lihrium  l.y  l.'.'.liiiK  thtni  "ilh  relativ.'ly  lar-ic  amounts  ..f  carl.ohy- 
. Irate,  which  nets  liy  pr.iteetin'_'  the  in-.'t.'in  of  tli.'  Ii.mIv  fr.im  .li'^iiitet.'ra- 
tioli  (see  |.av'e  oTli.  Kv.'ii  with  a  .li.'l  .'Xc.ssiv.'ly  ri.-li  in  .■.illi'iliy.lrates 
that  no  mor.'  than  .-..x.'rs  tli.'  .•al.ni.'  ie<|uir.'ni.'nls  of  tli.'  pati.'iif,  nilr.itr- 
onous  e.piilil.iimii  has  also  I..M'Ii  attaiiie.l.  The  pr.itein  minimum  \» 
which  I'.'ver  I'atienis  .-an  \n-  re.lu.'.'.l  is  ne\  .'ith.'less  c..nsi.lcral>ly  hi'4:her 
than   the   miiiimuni    in   n.iinial    in.lividuals. 

From  the  ah.ive  r.'Milts  as  a  w  h..l.'.  il  is  pr..l.al.ly  saf.'  t..  i-onelii.l."  that 
thei'e  is  n  siticljii  ili slnirlinii  «/'  pmli  m  •.'..ini;  .ni  in  tlw  li.xly  .luriin,'  fever, 
l-'urlher  .'\i.|enee  of  sii.-h  a  .l.'striicti.ni  is  furnish. mI  l.y  the  jtr.'seuee  in 
the  urine  of  e\.-.'ssive  amounts  of  .'reatinin,  of  imriiie  liases,  ali.l,  it  is 
sai.l,  of  ineoinpl.'l.'ly  hy.iroly/.'.l  pr.ileins,  siidi  as  the  allnimos.'s  ^  pr.i- 
te.is.'s.^  .M.ii'.'.iver.  when  ill.'  f.'V.r  -mhleiily  ti'iniinali's  in  crisis,  then- 
is  a  mark.'.l   in.'i.'as.'  in   the   ex.Mi'li f  urea    'th.'  .'pi.-riii.'al   ur.;i    in- 

crensei.  whi.-h  in.li.-at.'s  that   an  exi.'iisive  .l.'aminali.m  of  iirotein  Iniil.l- 
ini;  stones   (amino  aeidsi    is  .ici-iirriiiL'.      The  s.i-.'alle.l   "iliazo  rea.'tion 
olitaino.l  in  tli.'  urine  .lurinji'  the  iVv.'i-  is  aU.i  licliev.'.l  t.>  .leji.'U.l  on  the 
pr.'s.'U.'.'  .if  alinormal   prolein-.lislnti-uiat  ion    pi.i.lu.'ts. 

.\s  t.i  the  spe.-ilie  .'aiise  of  the  in.-r.'as.'.i  pr.il.'iii  .lisintevral  i.ni.  litth' 
is  known.  Several  factors  may  op.'iat.':  1  '  th.'  partial  starvation  ..f  th.' 
iiatieiit.  eritailinii  an  in.Teas.n!  l.v.'ak.jown  ..f  protein  to  ui.'et  the  .•ah.rie 
re.iuirements:   .  2 '    the  hiirli   t.'nip.rature.   \^lii.-h   in   its.'lf  may  stimulat.' 


i    i  i 


•    I: 


hi     m 


iiierensed  ))r.itein  meta 


h.ilisni,   for  it   has  ln'cn  shown  that,  wh.'ii   norii 


T'-M 


Ml   I'Mlcil.ISM 


1      • 


\        r  I 


I  u;lin, 


.iiiiliiaN  iii'r  ai  tiliri;ill\  uaiiiml.  iuhIimu  iinlal"iliMii  lici'oiiii's  iucrcascil 
.iiiil  lii  inxii-  iniitfiu  ilerciiiiixisilinii  |)n)ilii('ts  s|n'citifa!ly  i-aiisiiin  ati  <\ 
I'fssiM'  l)r(ak<lii\' u  111'  i)rii|(iii. 

Altliniiu'li  tiici'c  is  iiiiMcnscd  jiroti'iii  )>i-('aU(l(>\\  n  (luriii>;  fcvci',  it  imi>' 
imt  til'  fniiruttiii  tliat  Hilly  alinilt  'JH  Jior  cclit  cil'  the  lotal  I'Xlioliditiii  ■ 
..r  tli<'  liiiily  is  (l''ii\i'il  riniii  lliis  luoilstulT.  S(i  per  (Tilt  cciiniiiK  t'rum  imn 

iiili-ujii'iKMis  iiiatcTJal.   wliii'li    iiiiisl    l)p    fat,   1 aiisi'   tlic   availaMc   caili'i 

IimIiiiIcs  at'p  used  lip  at  an  oarly  staiio. 

Sii till-  Lii'iii'iai  iiii'taliiilisiii  is  ini-icasccl,  tlic  ixcossIm-  ln'caUdnu  ii  '<( 

tlic  t'att,\-  siilislaiiri's.  DiTiiiriiif:  as  if  ihns  in  tlio  in'cscncc  of  ii  diininislnd 
(■(iiiil)nstiiiii  (it' cailHiliydiatc's,  inlcircrcs  witli  tlio  pnipcr  oxidatidii  of  lli'' 
lattv  acid  iiiolcciilcs  and  leads  to  tlic  appearance  of  so-('alle(l  acidosis 
products  ill  the  urine,  and  consecpicnily  to  a  relative  iiicioasc^  in  tie 
urinary  aniiiioiiia  i  pa-re  tllti  .  A  li  mh  m  it  In  iii  Idu-il-i  tlicrel'ore  exists 
■|"iie  aeidnsis  may  reacli  a  consi.leralile  dcirrcc  ((f  severity  and  cause  llic 

tension  of  carl(oii  dioxide  in  tlic  alveolar  air  to  liccoine  diiiiiiiislicd.     Si 

a  similar  dcirree  of  acidosis  iiia>  lie  itrodiiccd  in  partially  starve(l  ani 
iiials  liy  oxcrlicatiiii:  tlieiii  witli  iiioisi   air.  lull   not   so  if  the  animals  are 

lilierally    t'c(l    witli   carlioliydrales,   it    is   pr(ilpal)l>'   safe   t nicludo   tlial 

aliniidaiic(    of  earlmlix drate  is  advisalile  in  the  food  tliat  is  fiirnislicd  to 
rev(M'  i>aticnts. 

Anotlicr   iiilercstiiiii   iiictaliolic  cliaiiire   in   fc\er  concerns  llir  .tnll   hti! 
iiiifi .    Tliis  In  si\ii|i"(|  li\  oliserviii'^  tlie  amount  of  sodium  chloride  (>xcrctcd 
li\    the  iiriiMv     As  is  well  known,  this  liecomes  iiiarkc(lly  diminished  until 
the  crisiv  of  the  fever,  when  it  siiddciil.N   increases.     Salt  retention  is  more 
marked  in  certain  t.\  pes  of  fever  than   in  others,  and  it  is  essontially  dif 
fcrent  in  nature  from  the  salt   retention  that  has  been  oliscrved  to  occur 
ill  iie|iliritis.     This  difl'ercnce  has  hceii  lirolludit  to  liplit   liy  pxaiiiiiiation 
iif  the  (diloiide  content  of  the  lilood.      Ill  lic|)hritis.  the  eoiieentiation  of 
r-hlorides  in   the  lilood   is  coiisideralily   increased,   whereas  in    fever   it    is 
niaikeilly  dimiiiished.     The  (leticienc\    in  salt   cliinination  cnil  not   he  at 
triliuteij  to  a  (lelicieiiev    of  salt   in  the  fond,  for  it  sets  in  liefore  the  diet 
has  lieen  curtailed  and.  when  salt   is  !.;i\-en  to  a   felirile  jiaticnt,  it   is  ic 
t.iiiied  in  the  lio(|,\    to  .1   urealer  dee-rec  than   is  the  case  in  the  iiornial 
iiiili\  iiliial.      l''or    some    reasiMi    the    tissues    in    fever    have    acipiircd    the 
|iioperty   of  rctainiiiir   \:\vj.v   ipiantities  of  s;dt. 

.\ttempts  to  study  lln    inilir  liiiiaini   diiriiii:  fever  have  frc(|Uently  lieeii 
made,  hut    the  te(diliical   diflieulties  of  sucli   invest iuations  make   the   re 
suits  iiiiceria!!!   .-'.nd   of  little   v.-i.lue.      Tha.t   some   retention  "f  water  (iccurs 
durine  fever  is,  however,  evidenced  liy  the  dilution  of  the  lilood.     At  the 
eiisis  this  hydremia  (piicklv   disappears  at  the  same  time  as  the  increased 


rONTKor.   OF    IKiKY    TKMI'KKATI  Kl      VMi    Kl  A  I  It  725 

.  rniiiiintinn  of  chltiridrs  is  uoing  on.     Chloiiil.'s  im-l  wiit.'V  \v'>ul.l  thno- 
H.ro  sc'cni  In  lirliavo  in  a  siniilnr  fasliion  <lniiiin  t'rvcr. 

The  Heat-regulating  Center 

In  ,ili  discus^inns  on  tlic  icirnlation  of  Innly  tcnipi^iatnro  aiul  i1m' 
ranscs  of  fi'viT.  it  is  assnmrd  lliiit  a  iicatrPLnilatinir  or  fliornioKrnic 
center  exists  soinewliere  in  the  brain.  It  is  Iiclievid  to  lie  locateil 
aliont  tlie  ojjtie  tlialanii  or  eorpora  striata,  for  it  lias  lieen  fonml  in 
laliltits  tlnit  (lestrnetion  of  tlic  liraiii  anterior  to  tiiis  re<ri"n  <Ioes  imt 
canse  any  elnmite  in  lioily  teniperalnre,  whereas  (lestr\ietion  liehiml  it 
is  fnjlowed  hy  an  entire  npset  in  the  heat  reiiulalinir  mechanism.  Fur 
Ihermore.  artificial  pnm-lnre  o*"  this  i)art  of  the  Inain  causes  maikcil 
elevation  in  liculy  temperature  in  raliliits  (heat  ll\Mll•tnre^.  Most  in 
tcrestiii'.'   experiiiiciils    have    licen    ivcoi'ilcil    liy    liarliour."    ulm   snccccilcil 

ill  apiilyinu'  heat  or  I'old  Ideally  i.i  the  reu'ion  of  tin ntcrs.     Ky  ihe 

application    of  cold,    increased    nniscular   mdalMilism,   on    tin'   "lie   hand. 

and   diminislied   heat    Inss.   nn    the   othei',    were   excited;    and    < \  clsclv  . 

\Wien  warmth  was  applied,  an  iiicrease.l  In'at  loss  and  a  diministiid  licai 
priicluction  were  nliserved.     Iriitation  "f  this  reiricm  ot'  the  lirain  in  nnni. 

as  ai'ti'r  eciel.ral   licmc.riliair<'.    is   also   tu mpanied    \>\    renmrkalde  dis 

lUrliai s  in  heat   reirulation.     It   is  lielicvcd  hy  many  that   the  essential 

canse  of  infections  fever  is  an  action  on  these  centers  liy  toxic  substances 
which  ilevelop  in  the  Mood, 

The  centers  may  also  he  acted  on  liy  various  ilrni/n,  some  of  which 
excite  them  to  inerease  the  liody  temperature,  others,  tn  lower  the  lem 
pcratnre  whoa  this  has  already  lieen  elevatcil.  When  snlntions  of  sodium 
chloride  are  injected  intiavennnsly  oi-  sidicntaneously  or  even  sometimes, 
particularly  in  children,  when  administered  hy  nnnith,  more  or  less  feviT 
iii;i\  result.  This  must  lie  a  speeilic  action  >if  the  Na  imi.  for.  if  iiisl-'id 
iif  pure  solutions  nf  yiiCI.  solutions  containintr  calcium  and  potassium 
>alts  as  well  as  those  of  sodium  are  injected,  no  t'ever  is  indm-ed.  This 
I'act.  taken  aloni:  with  the  cloi-e  siniilarily  lietween  |iuneture  dialietes 
.ind  heat  puncture,  lends  s\tiii)oit  ti>  the  view  tiiat  in  its  initial  staffes 
experimental  fever  of  this  typo  is  the  result  of  an  excessive  lireakdowii 
of  ■jlyeotren  in  the  liver.  It  must  not  lie  imagined,  hnwevcr.  that  ])crsist- 
iiit  fever  can  lie  attrilnited  to  Mwh  a  canse,  since  the  lever  remains  ;iftcr 
llie  .jlyccificn  h,i>  all  heen  rennived.  Other  chemical  sulistanees  ]irodnc- 
iinr  fe\er  are  caffeine,  certain  othei-  purines,  and  partii'ularly  tetra  hydio. 
iiaphthxlanun. 

Helonsrinir  t"  this   trrou!)  of   fevers  must   also   he  considered  the   im- 

|i(irtant  ones  prodii 1  hy  the  intravenous  injection  of  certain  forms  of 

protein,  as  those  of  cL'fi-  while  or  those  derived  from  the  liodies  of  liae- 


M 


) 


T2f< 


Mll\lliil.i.-M 


Ifiiii   iir    I'loin   till'   liiki'il   riii|iii>4'lfs   111'  ii    liiii'inii    hliMdl,      'I'lif    I'i'vir    in 

ihisi'     IMM'N     is     III!     lllllllll      CMllM'll      ll>      il      IMl'l'lllllliMII     cliwclx       irllllcil      111      lll.ll 

ii's|MMi^llil<'  I'll-  iiiiimlislii\i>   I  MM'  iiiiuc  Ml   .      Sii.'li   inj('.tioii>  i|<i  mil    im. 

diii-i'   I'l'M'i'   ill   iiiiihiJilN  iillt'i-  ili\isi 'I'  lln'  I't'i'vii'iil   >i>iiiiil    i-iinl  iir  i'\ 

<M>i<m  III'  till'  liiiillrijiin  It  i--  l>i'lii'\  cd  thai  liiiili>  i-ilx'S  iil'  Mi-riillnl  ii>i'|i 
fit'  t't'vcr,  ipi'i'iiiiinir  iillrr  M'\rii'  I'lUilUNiiiiis  nr  ciIIht  wniiiiils,  iiiiix  In- 
tin'  I'l'-llll  I'l'  (|i-strili'liiMi  111'  |ii'iili'ilis  within  tlii'  Imm|\  .  Siiiiiliiil.v  tin'  li-' 
ji  ti'iiiiii'i-atiii  I'  iliii  iii'j  iiit'i'i'li<>ii>  iiiiiv  li.'  ii\(  iliif  111  thi'  In  i-aUiluw  ii  |inili'i' 
111'  ihr   lilirriiiir'^Mliisiii    ill    ll rlU. 


in;  III:  .i  iki!'! 


I  .: 


i   i- 


',  !  . 


1!  %  ' 


Significance  of  Fever  in  the  Organism 

It  is  ini|iiissilili'  at  iiri'scnt  In  stati'  il('liiiiti'l\  wlii'thcr  I'l'vrr  is  a  ii' 
artiiiii  III'  till'  Di'iiaiiislii  auaiiist  sitnii'  iiirri'tiim  ami  tlii'icrnrc  nf  lirnt'lii 
ill  a'.sistiii!.'  till'  iii-«aiiiMii  In  rmiiliat  it,  nr  wlirtliiT  it  is  in  itself  an  iiii 
I'MMiialili'  i-umlitiiiii  'I'Ik'  iini'stinii  ran  rntainlx  imt  In'  aiiswci'cd  liv 
iilisiTvinir  I  111'  lirha\iiir  nl'  liartnia  urnwiim-  at  ilitl'i'iciit  tcniiKMaluri's 
in  vaii'iiis  Mii'ilia  iintsiijf  tin-  limlv.  That  I'lTlain  hai'tt'i-ia  slnnilcl  In- 
t'liuinl  nut  111  ihi-iM'  at  im-iiliatur  ti'iii|ii'iatni'i's  cinial  In  tlmsi'  I'imukI  in 
till'  Imily  iliiiiii':  t'l'vcr,  ilms  imt  at  all  pinvt'  that  this  I'i'Mt  is  nf  si'_' 
iiilii'aiii'i'  as  a   niraiis  nl'  riinilialinir   thr   lmhwIIi   nl'  thf  liartmia    in   tin 

lii.ily.      ll    is  iiiiiliinlitnl   that,   ulici-i'  tlii'  linily  ti'iii|u'iatiii'o  In im's  cn 

i-i'ssiv(l\-  hiuli.  tilt'  (•iini'i-t  li-i'atnii'iit  is  to  k»'i'|i  it  ihiwii  as  niiirh  a> 
piissililc.  Oil  tin-  iitlnr  liami,  lln'  ii'iliu'i'il  iiiiir1alit\-  that  has  t'nllnwi'il 
the  inlriiilnrliiiii  nl'  tin'  i-nlilliatii  tri'uliiii'iit  in  typliniil  I'i'Mt  iiia.v  mn 
III'  iliii'  sii  iiiiirli  In  till'  rcilnctinii  ill  Imily  trni|i('raturt'  itsclt'  as  In 
the  I'avcii-alih'  cll'i'i-t  |iioilm'i'il  nn  tlir  ih'I-miIis  systi'iii  ami  riiciilatinn. 
\Vt'  i'('i-taiiil\-  kiinw  that  in  nninial  animals  iiinili'iatt'  ilt'Ki't'cs  nt'  li\|)ii- 
li.vi'cxia  iirniliifi'il  li.\  ('Npiisuic  tn  iiinist  heat  art'  wi'll  Innni'  t'nr  i-iiiisith'r 
alilc  |M  ricitls  111'  tiiiii'.  thus  imiiratiiiu:  that  it  is  llit'  inl'i'i-tinn  ami  m>t  tin 
li\  liiTllii'i'iiiia  that   i-aiisi's  tin-  si'iiiius  ilaiiiau:i'  to  tin'  Imtly  in   intii't  inii- 

I'.'Vi'l's. 

MKT.Mit  iMS.M    ({KI'HK'KNCKS 

I  Miimiuraplis  ami  (iiiuiiial   I'aptTsi 

il.iisk.  (ii-;i)i;iiii:     TIm    i:i,ii,irils  cf  lln-  S,-ii' c.f  Nutriliuii.  \V.   H.  SiiiiiMJns  fn.,  i  .1. 

:;.  I  ill  7. 

•I -Mthi-.-il !,    I..     I'.:        rill-     I'lix -ii'liiL',*     "'     I'll  I'  ill     \li  !;il.(.li~iii.     \l •l;i.-i|>!i~    i.n     l;i.. 

.liriiiisl  I  v.   1  .t'ii-iii;iti -.  lirtt  II  \  I -)».,   l;ilL'. 
■|;ivl..'-.  A.   K.:      |ii-.-Mi..ii  :ni<l    Mi-IjiIiuIImii.  1..;,   \    |-,lii;;ri-,   V.  \\    \'i'\V,    I'.UL'. 
ifiirli  iliill,  F.  1'.:      Till-   l'li\>in|..^v  iif  thr  Aniiii.i  A.-i.ls.  Y:ilr  l'H•^^.  Nfvv    llinrii.   l;il.. 
"Mnflcn.l,  .1.  ,1.  l;.:      Iliiil.ciVs,  Its'l'iil  li..lo(;ir;il  Pliv<i..lf,i;v,  K.   Anu.M.   llH'!. 

■iFiirlli.  viii:      Till-   rMii.ii-iiis  nT   I'in  .i..l.i;;i.  ji  ;iii<i    l':itliiiliiuii-:il   ''I ^Ii,>.  •'•'■.  •'•   '•. 

I.i;i|>inc-utt   I'...,   I'.ilil. 
'I'.Tnncs.   W.:      \ii.-K  i.-    Ai-i.N.   MniiciLrrMiili-   in    lii.irli,iiii^ti-\-.    I.nii'.'iinni-^.  <;r'''-ii   *■   "'"■• 

lull. 

■'.M.li.l.l.    l.;ir;,u  Mr    r,,:       i;i:;rl.iii-M    .1.  l     I'iiv -mln^ir.    llill. 


m. 


(ONTKOI.   OK    honV     IIMI'IKAII  III     AM'    I- 1  M  II 


727 


'Liiilli.!-.  -1.   II.:     Tli>'   I'lii".   Mcu.iuiHI'l'"'  '»   "''"' !'""'>■•   l-"»;;"'»ji".  "'"'ii   1^  '■" 

\liilli.v\«.  A.  1'.;     l'lnMi.lMi{iiMl  rii.ini«tiy.  Wm.  \V I  \  i  ,,..  KM:. 

'hiikiii.   II.   K.:      (txi.liilh.il.  iiii.l   «.■  lurti.iiM  ill   111.'   Aiiiniiil   r."'lv.   .\|..ii..^rii|'li-.  m    Hi" 

1  liiiiii»trv,  l.iiii:;iiuiiiH,  (ill. 11  Jk   '■....   mil'. 
^I.iiilli...,  ,1.  it.:     I'ri.l.l.  iii«  ill  Aiiiiiiil  \l,liil.i.li»ni.  \'.»»'. 

•Kii  lt..i»,  K.  K..  an.!  .i.lliil i"!--:     di nl  i  ■|,..Mii«li  v .  I';i|..i-  I  i..  'J.V    V'.li,  li.i.  M.  ■!  . 

IIH".  17.  XM  xi\. 
T|lcl»'.lii-f,   y.  (i.:      Am.  .I..iii-    I'livM..!..   I'.'li.,   xli.  -'T.'.  ;iii.l  •^•'■:. 

-M.ii.l.'l.   I.iili.v.n.-   It.;      Il.in.y   I..  •■ ,  -I     II.   I.i|.|.iii.  .'tl   '■•■.,    I'.'H  ll'l'..  y.   I'M. 

■M.Ci.lliiiii,    K.    V I   ...lliil".riili.r-:      Num.  i..ii»    [.iipiM   ii.   .Iinir.    Iliiil.   Chrm,.   I..' 

uiiiiiiii;:  I'.'l.i. 

ILipkiiiy.   K.   lM.«i:iii.l.  nil. I   Will.  ...  k.   K.   '..:      .I..it.   I'l.>-i..l..    I!""-..   \vx\.   ■>*. 

"Mavli'.s,  W.    M.:     Th.-   n,vM..l.....v  <.(  I' I  :.ii'l    I.... in    hi   Ki.t.   I,i.ii»'iiiiiii«.  lir.'.ii 

&  (■...,   I'.'IT. 
i-M. Ciilluiii.   H.   \.:      lliin.N    l.ii  111...  .Ii.m.  Am.    \l..l.   .\»ii,.  Inl.. 

iSh.tI.  .1.    K.,  ('ill Wliiti'.    !•:.    1'..  nil. I   Niv '.     •'.:       '".II-.    I'i"l     <  li.iii..    I'.il.l.    XV. 

ISl;  iliiil..  I'.'l.-.  \xi.  :'.ii'.'. 
".Htc|.|i,  \V.:      Itliiilii'm.  /Isilir.,   l!"".'.  \xii.   I-".'-'. 
I    I'liiik.  ('M«iinir:      i;it'.l.iii>«r  ilrr  I'liN  «ii.|i.i;i.-,   I'.'l'.. 
i.M..Kill..|..     M.:        I' I     Viilm-:       \Mk.I     T1..-:      \<r    mi-I     II-u     '..    r;il..ihil.'    Tl...i... 

IJllllr.ll'r. 

I '..MfCiiy.  I>.   Majiir:     Tlir  I'lutviii   l-.l.iii.  iil    in   .Niiliili K.    \rii..l.l.  I I..11.   IHI-. 

iM'niilirVy,   Nf.  S.:     riii'mi-irv    ..f  l!.-|.ii:iti in  s.li.-ir,  r '..  T.  xt    lt....lv  ..i    I'li.vsii.li.u'.v. 

Isiis.  i. 

".Villi,  1".  r.:     (llyii.«iiri;i  iiii.l  l>i;il.i  I.-.  Hi.-l H'l  '•. 

1 '.lixliii:      lliiilirtrs.  ...  ,    ,- 

^'WimiiVHlt.   If.  T.,  Siinsiim.  W.   IL.  :iii.|   W  iLI.  i.   1.'     M.:       '"m.   Am.   M.'.l.   A.«ii..   I'.'l.. 

I'x   ,   JOI.r.      AUii   Tii>|..i.    A.    K,.   iin.l    Unit. .11.    I'.;      .I..ui.    I!i..l.   l-|..ni,,    ll'Ki.   \x\. 

-iMHil.iii'l,  .1.  .r.  1!.,  nii.l  I'nlk.  M.  K.:     Am.  .Iinir.  I'l  '     l!il7.  \lii,  I''-. 

--Iliininmii.  I...  nn.l   llii>.lil.;iniii:      Ai.li.   Inl.  M.'.l..  1  ..  Ml  7hh. 

j;(;,iii \V.    Ii.:      It. ..lily   'liiinv.'^    i"    l':riii.    Ilnnv"!'         -.r  I'H'l    '''"-"'•   ''■   A|.|.l.t..ii   A 

('11.,    I'.il.l. 
-'Knowltnii.  !■'.  r..  iin.l  Slnilini:.  1;.  11.:     .Ii.in .  I'li:  «ii.l.,  I'.Hl'.  xlv  .   1  1'.. 
.■I'iittiisi.ii.  S.   W..  nn.l   St.iilinu.    K.   U.:      .l-ui.   I'livsii.j..    I'.'l.:.   xUii.    l:-.:   :'l-"  <  '"I'l^ 

sliiiiili  ami   l':ili.  TM.ti:      ll.iil.,  i'.   II'.. 
-■■Mailri..!.  -I. -I.  I.'.:     (llyinlysis,  .Iniir.  Itii.l.  (  l!.  in.,   I'.'l:.  xv.   I''.. 
-;\lnrliii.  .1.   1;.:     -li."!-.  I'inl.  CIkmii..  H'K!,  x\  i,  7'.'. 
--Criiii  ksh.-iiik:     .li.ir.  I'l  v«i.i|..  l'.il:i.  \h  ii,   1. 
-"Mai'lfo.l.  .1.  .1.   1;.  tiii'l   I'.iiri-i',   l>".   (i.:      Zrlitrall.!     I'.    I'liy-i..l..    I'.'l::.   xxm.    1-1 1. 

;..\V halt,   H.  T.:      .Ii.nr.   Am.   Mi.l.   A>«ii..   I'.'l''..  Iw  i,    ll'H' 

HViin  Slvkr,  1).  H.:     Tin'  I'ri'Miit  Siyniti.-.'iii -i'  nt'  tin'  Aiiiiii..  .\.  mU  m   I'l.v  M..li.;.'y  anl 

I'.-ltlml llaiM'v    LntiiiTs.   .1.    I'..    l.i|.|.iii.-..tt    \    ''.■..    l:'l.'.  Il'l''..    |..     I  I'''.      Al>.. 

).a|..Ts  iii'.li.iir.  liii-l.  flirni..  I'.'l  I.  ix.  Iv",:  xii.  J7.-.:  il.i.l..   M'l-.  xn,  ::i>l  :ni.l  .;'.'■': 

iiii.l.,   l!'l",  xiii.    1-1.   1-"'  ■■"I'l    l^'-  ,     ,      ,    ,  .  ,  .         I 

•-F..liii.  (I.,  ami  l'.ni-.  W  . :      .I...1..   I'.i.  I.  ''Ihih..  xi.  ^7  1    l'.''. ;   il'i'L.   I'l-.  ml   "  :""l 

:;.■■■:!. 

•Xl.i'l,  .1.  .1.:     Til.'  .\l.  Il..n  Li'ilnrr.  >ri.  ill.  .   1'.  I.'..  ^Iii.  1  '•  '  .    . 

:;.il,.»l,.tt,  A.  W..  dill..  It.  1..  (>.,  Wi.kri;.  A.   !>.:      Ar.li.   1..'.   .M.'l..   H'l''.  nmh.  ' 

:^-'l.»s...'.  .1.   It.,  aii.l   Vail  Slyk,',  D.  !>.:     .I.mr.   A....   M.'l.   A-.mi..   I'.'l  7.  .  l.ii.  '.'I. 

^'■•SlialT.'i',   1'.  .\.:      Am.  -Ilim'.   I'livsi.il..   1!'"^.  xxxiii,    1. 

-Catli.art,  K.  1'.:     .Imn-.  I'liyi-inl.,  I!"i7,  x\\v,  ."im. 
:>Mvi'rs  ami  Kin.':     .Ii.m.  ni"l.  Clii'm..  llH::.  xi\.  H. 

::.|,.\i'ii.',  r.  A.:     rf.  W.  .[..Ill's. 1 

'".Imiu'S.   \V.:      Nnili'i.'    A.  i.l~.    M ^liil'li-  •'"    I'.i...  In  ii;i-ti  y.    l..'nym:M.-.   (.i.'.'i    .V    <  •'■. 

lull. 

"  11.11.'. lii't.  S.    Ii.:      IliiiA.v    1...  tur.'.    r.Ti  H'. 

i-liiint.r.  A.,  anil  (iiv.iis,  M.  11.:     .Ii.ur.  llinl    «lniii..  I'-'H,  x^n.    •"•••• 

'  Huriaii    li     an.l  S.liiir,  II.:     Cf.  Ma'-lcoil  in  It.'iinl  Ailvam-.'s  m  l'liy>i..l..j.'v  :ia.t  Hn. 

'.■I,",ii.i"-trv.  ..i.  hv  I.i'c.iiar.i  llil't,  V..   Arn.-.M,  I.ai..!nii.  !!»".'. 
"M.'n.U'l,  Ijifav.'tt.'  It.,  ami  I.ymnn.  A.  V.:     .I'mr.  Iti.il.  Cliiii)..   ll'l".  mm.    M-'. 
■     i,|  K.is,.    \V.  r.:      .1.'   1.  !!i"l.  clii'in..  tl'l-"'.  n".   H'-'- 


■Tavl.ir.  A.  K., 
'll"|ikiiis  K.  «; 


.1   11..,.. 


W.    I! 


.I..UI 


I'll 


1.1. .    Is'.'!' 


i    i 


w\ 

1 

I 

t 
,1 

1  I 

i       '^1      ! 


.hi 


(•»»8  MKTAWH.IKM 

•  ;\...li.  M,  iin.l  Itvr.  0,:     ZDh'lit.  l.  I'li».i"l.  I'h.in,  \\'v\>,  hm,  :C\>;  il-i'l,  lull.  Ixm 

.11 '.I. 
i-M.CIiirf.  (!,  W..  ViiKTiil,  II.,  nn<l  I'latt,  .1.  Hi      Am.  .I.hii.  I'liynii.l.,  llHtl,  xlii,  5UI1. 

•  ••MI.Kir,  \V.  ».;     .I.iiir.  Mini.  Chrm.,  IHIJ,  »i,  ll".l;  i\M..  Ililil,  xv.  IM;  il.l.l.,  IIMI,  »m. 

.-ilT;   ilii.l..    li'IJ,  xi,   HI;   il.ld.,    I'.'l.".,  %\i.    l:'l;    iI'LI.,    lOlt,   xix,   I;   ililil.,    IKl."., 

\slii,  :ilT;  jMil.,  Il'lt.  x\ii,  :iiri  iliiii,  liil.'i,  »xii,  l:i.'t.     Al»i(  lllo.it  ninl  Kiiiiij 

.I.Mir.   Itiol.  ni.'iii.,   Iltni,  xxvil,   1117;   il>i,l.,    lUlii.  xxiv,   »I7;    IMmir,  .Icwiiii  mil 
ll.irti.r:        II. i. I.,   ll'lil,  xxvi,  417;  il'i.l.,   H'lU,  xxv,  :>". 
••  I..  !illi.«,  .1.   ».;      Til.'    r:it<,  Mnn(iurii|'li«  i.ti    Iti.i.  Iii'ml»lry,   I...ni(iiiiiii«.  (Iri.n    .1    '" 

.(■.,., |,...  K I   \|..ll.;.i...  \     II.;      .I..iir    l'ln.i..l  .   I;>1  t,  \li>.  .''.:   iLhi  ,   ll'l."..   xiix,   I '.7. 

i;i.|..r.  II.  S.:     .I...I   .   III..!    (  1„  II,  ,  I'.il.:,  xi\,   117. 

■  >liii..|l.'V.    I.    I». :      I'r.".    I'll.*-     S,.,'..   .Ii.ui.    I'liv-i..!.,    ll'l;.',   xl»,   J".. 

•  Ilill,  I."  ..ii:ir.l:      .\.Mrr-i.  I..  Ihc  I'livs.  >..'.   I'.iit.  .\.<»ii.  for  th.'  A.h.  of  .McL,  SimIi.hi. 
■r,  I'.ilJ. 

■  HImir.r,  I'.  .\.,  iin.l  (•..l.iimii.  \V, :      .\r.li.   lul.   M."l..   H""'.  i».  '>■■'■•*. 

■••Iljirl..iiii,  II.  (i.:      Ai.h  I'.   i:\|..r.  I'litli.  u.  I'liaii. .».•..   li'TJ.  Ux.   1.     Al-.i  DiirlM.ur  inM 

Will;;,  S.  M. :     .I.nir.  riiritni!!.'.  nn.l  lixp.r.  'rii.riii..,  l!'l::,  \.  in.-.. 
■;|l.«l.'tl,  .v.  W.:       -!..n. -iiipl'i.-   M.  .ii.iii.'.  I'.    \|.|.l.'..ii  A   r...,  1!I17,  i. 


.*     'll 


'I.      . 


V:\\{\    VIII 
TIIK  KNDOCKINK  ()K(iANS.  OH  DKTLKSS  (ILANDS 


CHAI'TKU  l,\\\l 
TIIK  KMXM'UINK  <>|{(;.\\s   (tU  l>l  (TI.KSS  (il,A\l>S 

III  oi.ln-  tlml  the  vaiious  niMivilics  of  tiio  niiiiiinl  <.rKiini<.iii  iimy  u.'» 
.m.'iPiitly  ns  n  ^liolf.  if  is  lu-wssiiry  fliat  iIm-s.^  oC  oiip  pint  !■.•  rorrflatci 
uith  those  of  aiiotli.T.  This  correlatioii  of  fiiiirtioi,  is  iii.Miialnl  rith.'r 
throiij.'''  tiic  nervous  system  or  throiijili  the  act  ion  '  n  "ne  part  of  the 
i.o.ly  1. .  .iistnnees  i.io.liiee,!  in  aiiotlier  part  aii.l  earried  t.etweeii  them  l.y 
the  l.h.  ,.  Control  thn.iiu'li  tlie  i,ei  \  ous  system  is  cspeeinliy  developed  foi- 
llmse  funetions  w'ieh  have  to  ).e  l.roimht  promptly  into  play,  siieh  a- 
miiseiilnr  movement  and  th.'  other  physiolotfie  proeesses  .■oneeriied  in  the 
ndjustnii'iit  of  the  oriranlsm  to  .piicUly  ehnnuinR  eonditions  of  its  environ- 
mrnt.  Control  thionirh  thel.lo.id  is  the  moehanism  hy  whieh  the  metaholie 
aetivities  of  dilTorent  oilmus  are  mainly  correlate.',.  The  ehemieal  snh- 
sfaiiees  involved  are  often  called   iiilininl  .inritiotis. 

Some  of  those  internal  secretions  are  merely  hy-products  of  metabolism, 
and  are  only  incidentally  ns.'.l  f..r  the  i.nr)>ose  of  hrinfiin-.'  a')..ut  control 
between  different  parts  of  the  body.  To  this  irronp  beloiiL'  carbon  dioxide. 
«  hich  mny  act  on  the  respiratory  and  otlier  nerve  centers,  aii-l  nrea,  which 
amy  slimnlato  increased  activit.\  ><  the  Ui-lney^.  Indeed,  the  list  of  snb- 
stances  in.dndfd  under  sii<-li  a  delinition  of  internal  secretions  is  almo^' 
illimital)]e,  and  to  designate  by  the  special  name  of  lioniione  every  eon 
stituent  that  can  alTect  physi(do>.'ic  t'nnetions,  as  sonn-  have  ilono,  can  lead 
only  to  confusion.  The  intet-nal  secretions  with  which  we  are  more 
directly  concerned  are  those  that  are  specially  produced  for  the  l)urposc 
of  coiitrollinsr  the  metabolic  fniictioiis.    They  are  i.'iven  the  jioncral  naim' 

nf  anta.'oids  i  K.  A.  Schal'en.'       .\iiti i<ls  iiia.v   I ither  the  sole  •  riMluct 

of  sorao  special  uland  or  a  secondary  jn'oduet  of  irlands  which  hn      othe: 
functions.    To  the  former  class  belon-.'  the  autacoids  produced  by  t..e  pa-a 
thyroid,  thyroid,  pituitary  and  adrenal  irlands,  and  to  the  latter,  those 
produced  by  the  jiancreas  and  tfcnerative  (jlands. 

Autacoids  have  further  been  subdivided  by  Schiifcr  into  two  classes 

7'jn 


I 

'i 


m  '  t 


% 


■  i^l 


* 

'^'['^ 

1? 

M  1     ' 

■  \ 

<'.\  •  H 


"I'li; 


il 


,    I 


!     1 


i         ! 


730 


Tin;  i.mmxhim;  kkuans,  i>h  in  i  ti,i;ss  (ii.a.nds 


jiccoiilini:  ti)   wlii'tlicr  they  i\cito  iiH'liilmlii'   iiroccsscs  or  ilcpi'css  tlii'iii 
I-lxMiiiplis  111'  i\(itiil(ir\    iiiitiic'iicls,  ,ils<i  (Icsisjiiatcil   iis  Idiniiiiins.  iiii'  tin 
r|iiiii'liliiilic  |>lH.|ii..,.l  \>\    till'  ■ulrcniil   irlilllds,  wliicli  excites  llu-  liTlliiliJl 
ticiiis  ol'  tlie  >,>  iiip.itlielie  iieiMiiis  sxsleni,  iiml  i)ituitriH   iiiiiiliieed  by  111'' 
|i,,Nteii(ir  l.ilie  of  the  i.illiitiiiy  ul.iiid,  uliiidi  excites  l>l;iiii   iiiiiseiiliir  liher 

Iniiil'itiiiii  ,-iut;icc)i(ls,  mIso  cjillerl  .'/<(/'<//«  v.  iire  not  sue ininily  known.  Imt 

•■ii-e    ilinstriited    liy    tlie   sulistiince   ciiiit;iined    in    extract    nl'   tlii'    ])liieenta. 

\\lliell    len<ls   tii    ]ll'e\ent    tile    seiM'etilin    ul'   lllill<. 

AiitacniiU  nia\  lia\  >■  either  an  iniineijiale  or  ;i  i|ela>  ei|  act  inn  ;  liie  ellect 
wliic-ii  tiie>  |.ni<lnce  may  lie  liUe  tliat  witli  wtiieli  we  are  raiiiiliar  as  tin' 
lesnlt  nl'  stininlatidn  ul'  tin'  nerve  siipi'l^  "I'  a  uland.  lieinir  illiistraled 
aiiain  li\    the  elTect   nl'  e|iine|iliiine,  cir  tliey   may  act   sd  slowly  that    it    i-- 

,,iil_\-  after  a   vidcMaMe   |ieriod   i.l'  time  ilnriin:   wliieli   they   liave   lieeii 

in  the  ni-eanisiii  in  excess,  ih.it  an>  a|i|iarent  el't'ect  is  prodlleed.  Tli' 
shiw  ly  acting  antacuiils  Inne  lieci  i-aMccj  iii(iri>liiiiii  m  In-,  and  tlu-y  ai'c 
well  illnstrated  in  the  internal  secn'tions  nl'  the  anterior  lolie  of  the 
jiitiiitary  and  of  the  'generative  udainK     secretions  whicli  atTeel    <_'rowtli 

Hc'j-ardinv  lln  cIk  iniral  milun  n(  miliuuiih.  i-ertain  facts  stand  ont 
lir.iliiincntlv  .  licinu-  very  lar'jcly  the  |n"'ln.-ls  ol'  '.elands,  it  inisjlit  lie 
inn-iLrinc'l  Ihal  the\  wonlij  lii'  en/vinii'  in  natui-c.  I'>ir  en/ytiies  arc  iio\\ 
known  t.i  he  the  nnisl  iiii|iiirlant  active  ayciits  in  liioiilasiii  as  well  as  tin 
a'-ti\e  agents  in   iiian\    "I'  the  external   secretions.   liUi'  those  of  the  sali 

viiry.  L'aslri'-  anil  intestinal  iilands.     .\nti iiK.  imwcvr.  aiv  not  ciizynu- 

'I'hex  are  far  sim|iler  in  chemical  stinclnrc,  and  are  not  ilesti'oycd  !i.\ 
heat  in  the  presence  of  water.  The.\  arc  represented  li.v  a  eomparativel.^ 
small  molecule,  and  are  therefore  dialy/alilc.  Tiiis  latter  I'act  jiistifie- 
the  hope  that  it  ina.v  he  possilile  to  pre|iare  them  oi'  their  simpler  siilt^ 
in  crvstalline  I'orm      a  hope  which  lias  alreaily  lieen   ri'ali/<'d  in  the  case 

of  at   least   ol f  them      epinephrine,     tireat    prnyress  has  likewise  hee? 

made  in  isolatiii'j  the  a<-ti\e  principles  of  the  th.vroid  and  of  the  alitorim 
and  piisleriiif  hilies  of  tin-  pitnitary  ijlamls.  Tn  sniii  uji,  then,  we  m;\} 
sa.v  that  an  autacnid  is  a  spi'cilic  oryanic  siilistance.  formed  hy  the  eolN 
iif  one  uiLian  and  secreted  into  the  cireiilatiiiLr  tinid,  which  ci  ries  it  t" 
other  oriiaiis,  npoii  which  it   produces  el'l'ecls  similar  to  those  driiirs. 

Methods  of  Investigation 

'I'll  iii\esli'.iate  ihe  function  of  iiii  anlaenid,  careful  s1udie^  are  made  "f 
ihe  effects  pi'iiluce'l  I'  I'V  excisinn  "f  tile  Lrland  \\  hich  furnishes  lie 
autacnid  and  i 'J  li.\  adniinisterin'j:  int  ra\ '■in'Usl.v  or  sulicutaneoiisly  "i 
iM'alh  extracts  picparcd  frmii  the  iilainl  l-'reipiently,  also  liyht  is  thrown 
,,,j  the  fMi'cti'in  ,:>:  '111'  ;inl;iiMid  hy  i ihsc'v i nLT  the  elTect  which  f"l 
I'lws   pi-iiliiii'jed   ficdiii'j   v^itli   the  enili'i'riiic  uriraii  that    manufactures  it 


THi:    INIKKHlM.    (iKIiAN.-.    OK    l>l  (    II.K^    (il.WKS 


731 


i,„.l  \>y  olivTviii-  til.,  imllinln-ir  .•liiin-i-.  ill  til.,  vaiions  .■niLi.Tiiu-  ofgans 

"ill    .liSfilS.ll    .•.,ll.liti..ll^.        KlulM-Volnui.-    iin.l     iri^l..l..L'i.-    ^tU.li.■S    lllV    mIs..    of 

llii.   t;iTatcst    iiniM.rtau,....      A     '    ti.-ully    in    invest iirMtiii-,'   ll..'    fnn.-tion   ot 

nil  .•ii.l.KTiiic  or-Mii   lies   ill   111.'    f^'-t    llnil    ll."  MMT.^ti..!.   of   ii..  -iaii.l 

ii.'ts  iii<l.'|.cii.l.'iillv  of  tlioM'  rr.M.i  ntli.i-  uliiii.lv    On  111.'  .-.iitrary,  tlicrc  is 
un.|..iilit.M|ly    a    .■Iom'    .•i>-.,,..ial  l,,n    ol'    ruii.-l  i"i',   -"    ttiat    w.'    .'an    n..t    ti'll 


ulictli.T  a  .-lianu.'  <•(  run.'ri..ii  .'1.x  i\  .'.l  aft.T  ivinoval  ..T  s..iii.'  ulaii.l  of 
;i.linillistriiti..li    ..!'  ^'in.'    .■xtra.'t    i>   a   .lir.'.'l    .•i.iim'.|1i.'1i.-.'    ..I'   tlic    .'Xpfl'l- 

„„,„(;,1   |„-,„-.'.;iiic.  ..!•  i>  ill. In.'. '.1  liv    S..M1.'  X ii.ian    .'iT.'.'t   .l.'v.'l..|...(l   ..n 

MiM.lli.T  .■ii.|...-iiii.'  ..ii;ah.  It  uill  iin  .l..ulit  lak.'  iiiauy  y.ws  Ik'I'i.iv  siiT- 
ti.'icnt  .liila  liaM'  1..-<mi  .•..ll.'.-t.'.l  t..  .  iial.l.'  u>.  .l.'tiiiit.'ly  I.,  state  wliat  the 
|.ai'ti.'iilar  luiieti.ni  .iT  .a.-li  .•n.|...Tiii.- ..iL'aii  may  lie.  Siii.'c  iimst  iH'.eress 
lias  l.e.ii  iiia.l.'  in  .•.miieet i.in  with  the  a.lr.'iial  irlan.l.  it  will  !..■  a.l'an- 
lau'e.ius  t.i  e.iiisi.ler  the  fun. Mi. .lis  ..f  this  .jlan.l  first. 

ADRENAL  GLAND 

111  iiiaiiiiiials  the  a.lnnal  uhiU'l  is  e.iiii|>..se.l  ..f  tw..  parts,  the  .-..it.'x  an.I 
the  iiie.lulla.  In  ..tlier  er.ill|is  ..f  animals  Imwi'Ver.  th.sf  two  are  more  or 
l.'ss  sei)arate,  li.Miii.'  eomph'tely  so  in  fishes.     This  n..t   infre(|uent  sei)ani- 

lion  ..f  e..rte\   ami   <lulla   suee.'sts  a    .litr.'ivnt    fiin.'tioii    l..r   th.-   two 

stiii.-tni-es,     Kxperimeiital  iii\  .•sti'jalioii  supports  this  \i.'w. 

The  Cortex 

The  e.irlex  ..ii  ini.-r..s.-opi.-  .xamiiiati.in  is  s.'.'ii  to  1..'  .'.inip.iseil  of  ro\^s 
<if  epithelial  .■.■Us  arraii^cl  more  or  l.^ss  in  .■..lumiis  except  at  the 
]ieri;ihery,  where  tli.'v  form  ul.im.rnlar  niass.'s,  an.I  inxt  the  iiiedullil, 
whei'e  they  assume  a  reticular  formation.     The  cells  of  the  <rreat.'r  part 

..!'  the   cortex,    unlike   tin. si'   of   the   i lulla,   .■..ntain    no    '.jranul.'s    with 

special  staiuins:  .lualities,  Init  ih.\  .1..  .•.■iitain  parti. ■l.^s  wlii.-h  are  1..'- 
ii.'ve.l  t.i  li.'  comi.ose.l  of  .■ho|.'st.'r..|  esleis  an.I  Irritliin.  In  the  cells  ..f 
tne  reticular  porth.n  >'\  \\\v  .■.irt.N,  hou.'\.'r.  i.iuiiient  particles  are  n..t 
liifre.piently  oliser\.'.l.  Th.'  l.ioo.l  supply  ..f  the  cort.'X  is  ii..t  n.'arly  s.. 
rich  as  that  of  th.'  me. lulla,  li.'ini:  r.'pr.'s.'nl.'.l  l.y  fin.'  art.'riol.'s  w  hi.'li 
luii  inwards  from  the  cai)sul.'  t.iwar.ls  th.-  im .lulla  in  the  coimective  tis- 
siii'  that  li.'s  l.efw.'.'u  tli.'  .'oluiiins  ..f  .'orti.'al  .'cIls.  N.'i'ves  similarl.\- 
|..'lietrate  int..  file  .-.irt.'X,  s.nii.'  suppl\  in'.^  ils  1.1... ..I  vessels  an.I  c.'ll 
.'..lumiis.  hut    111. .si   of  th.'iii   pr...-iT.rni'j   t..  ll.e  ii.-,lnlla.     Tli.'.\    ar.-  .l.'rivi.i 

fr.mi  a  n.'tuork  of  ii.'r\.'  lihei's  in  tli.'  .-ap-nl ih.'  oe-aii.  an.I  th.'  n-'r\.' 

supply  of  this  n.'tuoi'k  .'oiii.'s  partl.v  fmm  th.'  suprai.'iial  i.lexus.  aii<l 
partly  from  th.'  splan.'hni.'  n.'r\.'.  l-liiihrx  .'h.iii.'a.llx  th."  .'.irt.x  is  .h - 
ve!..po.l   i"r<.iii   th.'   .-.'lis   ..f   th.'   u'cnital    ri.li-e,   that    is.    fnuii    meso.lermic 

.'.'lis. 


5     ' 


m 


a 


m  ! 


i\< 


I- 


■\ 


i      ^  !l 


i'ilHI 


;.  i 


782 


Till,    i.NIHM  IIIM,   nUiiWS.    (IK    1)14  TI.I>S   (Jl.ANKS 


\'iTy  littli'  is  kiiuwii  (•■iiii'fiiiiim  ll"  fiiiiclitin  itf  the  adrrnal  cortci. 
.lirlKMi^li  tlicic  is  little  (Idiilit  that  it  is  I'loscly  rolatcd  with  tlio  develop 
iiient  111'  llie  sexual  organs.     Tlie  ix  iilciu'e  for  this  is  as  follows:  (1)   in 

rases     nl'     seM|,-||      |MecMH-it\      il      is     I'dllllij     tliat      tile     ailrelial     eiifteX     is     lllUel 

hyperlropliicil  :  'J  it  heediiies  hyiiciiroiihied  durilij,'  i'i'e'J:ii.nney;  (3j  it 
is  ill  ill  veloi.eil  ill  sexiia'  deliciciK'v :  '4'  eliaiiv'fs  occur  in  if  during  tip 
estfiial  e\cle  ill  many  animals;  Tii  al'ter  castration  it  is  said  to  he  hyper 
trophied:  ili'  tin'  inneiiinist  )iortinn  ot'  the  cortex,  someliiiies  called  the 
hoiiiidary  zone,  is  much  liyperln.pliied  in  tin'  liiimaii  t'ctiis,  hut  this  hyper- 
trujihy  entirely  disappears  after  the  tirst  year  of  •■xtrauterine  life. 

Whether  the  cmtex  pdssesses  other  fiiiietioiis  is  diflicult  to  say.  Some 
tacts  uoiild  iihlicate  that  it  does.  lAir  examiile,  the  passable  of  hloo  1 
t|iroil'-'h  the  curlex  het'cife  reachiin^  the  medulla,  would  seem  1o  indieate 
that  some  chain."    wliich   is  jireparatory  tn  the  main  idiansrc  occMirrin^j  in 

the  iiiediilhi  tak.s  |)lace  in  the  hi I  \vhile  it  is  in  the  cortex.     This  view 

is  partly  suhstantiated  h\  the  ohserxatimi  that  when  an  excised  portion 
dl'  cdftcN  is  inciiliate<l  at  iiody  t eii!|ier,'it lire,  a  siihstaiicc  develoiis  in  il 
which  has  .-iii  aetinn  liUe  that  of  the  hormone  of  the  iiieiliilla  ejn 
ncplirine.  It  is  pus^ihle,  Imwever.  tlial  this  action  is  due  to  the  fact  that  ecr 
tain  ot'  the  deeumposit  imi  iirndnet»!  '<(  pnitein   ih'xclop  an   epinephrine 

like   aetinll       see    p.'l'^v   .""(l"-'  '. 

The  Medulla 

Ilistnlnuieally  the  liiediilla  is  eoiiiposed  "(  masses  of  polyLTonal  cell- 
with  hlo.id  sinuses  iiet\>(en   them.      The  hludd  siljtply  is  derived   from   vcs 

sels  that   have  i>r eded  to  the  mflnlla  Ihroiijrh  the  capsule,  and  it   i- 

cNtr.-mely  rich,  heiin:  inded  tlie  rii'hest  hlood  sujiiily  to  any  orpan  in  tin 
h..d\\  "jreati'i-  e\en  th.aii  that  t.i  the  thyroid  uland.  The  nei-ves  form  .i 
dense  ).lc\lis.  .■Ntcndiiii;-  intu  .•iiid  hetvM'cn  the  secretory  cells.  The  lin's' 
eliaiai-teristic  I'eatiire  nf  the  cells  i'oiiil)osin<;'  tin-  lueiliilla  is  the  presein-e 
ill  them  of  M-ranules  which  stain  readily  with  eliromie  aeid,  and  are  hence 
..t'teii  called  chrotiiiiifii  <-»//.v.  There  are  also  some  cells  coutainin'jr  coarser 
-raiiiilcs  that  arc  soliihle  in  water  and  do  not  stain  with  ehrome  salts 

Kmhi\ol..uicall\-  th.'  inedulki  is  devi^loped  frmii  the  same  neiirohla.stie 
■•ells  tliat  ffive  rise  to  the  sxmpathetie  nervous  system.    This  evidence  oi 

tl !,,se  ass..eiatinii   hetwieii   tlie   niedilllii   and    the  sympathetic   ner\iiii- 

system,   we  sh;ill   sei'  t..   he  suhstantiated   h,\    llie   results  of  experinieiita' 
in\  est  i'jation. 

On  aeeouiit  of  the  anatomic  relationships,  it  is  impo.ssihle  to  study  tie- 

.•iTeel    ..f  eXeisi..ri    "f   Iiie   CMrfex    ;n:d    laeihll!,!    s'-pai  a'el.\  .    or.    ill'lee^,,    e,    t.ie 

action    of  luire   extracts   jiiepared    from   either   of   these   portions  of  tin 


■nil    I MiiiruiNi    iil!(i\\s.  (ii;   nil-Ill-^  iii,\\i'>  <•'•• 

uliiiitl.  Our  iii\<sti!.'iiticiiis  must  ciiiiccin  liic  clVcct  ul'  iciiioviil  of  tin' 
whole  glaiKl  ov  ..f  tlif  iii.jo.-1i(.ii  ..r  .xtiiuMs  of  it;  Imt  as  wf  procncd  to 
..xamino  the  ilala,  it  uill  h.r„nu.  ..vidfiit  tliat  most  of  tli.'  cffc-ls  ob- 
served to  oceiir,  with  the  .NC'i.tiou  of  those  already  mcnti.iiiea  as 
d.'liTiitely  ass.ieiiilcd  wllh  the  rniticMl  liuict i. HI .  eiiii  li.'  nt t rihiil .'d  to 
llic  iiii'dulla. 


j,!, 

f 

1 


Adrenalectomy 

Kxrisioii  of  the  adrenal  uiaiid  in  most  Miiiiiials  is  very  quickly  fatal, 
the  only  well-kuowM  exception  l.eiiij.'  in  the  eMse  of  the  white  rat.  in  which 
excision  of  hoth  adrenals  may  not  1h.  incnmpalil.le  with  life.  For  smnc 
time  after  recovery  fmin  the  ancsilictic  tlie  animal  upon  which  doiil.h' 
;,dreiialectomy  has  l.een  performed  usually  h.^haves  in  a  perfedly  normal 
f^ishion.  althniitih  it  may  he  less  lively  and  less  inclined  lo  feed  than 
iiMial.  Very  soon,  however,  -,'eneraliy  within  twenty-four  or  forty- 
eisilit  hours,  delinite  symptnins  of  musciihir  weakness  are  apinirent.    Thi- 

wciikiiess  soon  hecoiiies  extreme,  and  is  :h nipanicd  hy   a    feel.le  jMiKc. 

a  depression  of  hody  temperature,  and.  later,  hy  .lyspnea.  After  an 
interval  Avhich  is  iicvim-  longer  than  a  leu  days,  '  h  supervnes,  heini: 
voiiietimes  preceded  hy  convulsions. 

When  only  one  adrenal   is  reiimve.!,  very   feu    „nimals  suceuiiih ;  and 
if  some   Unu-   is  alhiwe.l   to  elajise   so   that    the   inimediiite   shock   of  the 
,,peration  has  disat>peare.l.  it  will  usually  he  found  that   removal  of  the 
re.nainiiiir  adrenal,   althoiiLdi    ultimately   fatal,    is    not    s,.   .piickly   so   as 
wlieii  hoth  irlands  are  removed  at   oiie  operation.     The  reason  for  this 
lesiilt   is  that   o|.i.ortunity  i>  i-iven   for  a  compensatory  hyjiertropV  "I' 
acccssoi'y  adrenal  hodies  to  occur.     Su.-h  accessory  adrenal  bodies  may 
he  compose.l  of  cortical  or  medullary  tissue,  hut  it  is  tlie  latter  that  is  of 
iiMportaiice  in  the  ].res.-nt  eonneclinn.      rhromaftin  tissue  is  f.iuiid  in  most 
.inimals  aloii^'  the  front  of  the  aorta,  l.clueen  the   renal  arteries,  where  i« 
c-aii    usii.dly    he    reeotrni/ed    hy    slaininu'    the    tissue    with    .-hroiiru-    ai-id. 
Sometimes     ac'cssory      chromafliii     tissue     is    loeatcl     in     dist.-mt    parts, 
as   in   th.-   ei.ididymis   of   the    rat.    lor   example.      It    is   said    lliat   life   .-an 
he  maintained  if  oiie-eid.tli  of  the  total  amount  of  the  adrenal  suhstance 
he  present  in  the  hody.     Attemi.ts  to  prohmi.'  life  after  adrenalectomy 
hy  adrenal   transplantation   have   almost    invariably   met    with   ne<ratiye 
results,  because  the  (.'raft  nnderi:oes  a  raphl  process  of  necrosis  and  dis- 
appears; althou^di  it  is  said  that  1 1  aiispianlation  may  sometimes  be  suc- 
cessfully accomplished  if  the  ^'raftinur  is  done  into  the  kidney.     Admiiiis- 
M-ation'  of   suin-Mren-.l    extract    is    .-il^o    without    delinite    benefit    aftci- 
adrenalectomy. 


'      I 


f  r 


'i.i" 


TM 


rill    i\i>'iiuiM    ciniiw^.  Ill;  Ml  I  I'll  »  lii.wns 


^.^1 


I        ■  ! 


■  ■  .' ) 


if 

„  i      i      .     15 

I'     ''I 


»:        Bi 


!!M    :i 


^' 


i  i 


i  -ii 


yi 


i-i^ 


Suprarenal  Extracts— Preparation 

lii.ii'ctiiiM,  pjii'ticul.irly  iiiti.n ciiuiis.  u\'  extract  of  tlic  adifiiiil  >iliiU'i 
li.is  t'liniislicil  Us  with  must  (if  tlic  cvidciH'c  iipmi  wliu-li  mir  l;no\\  Idl'.'' 
rcmiriliiiji  till'  tViicliuii  <i|'  tliis  uru-.-ui  ili'iintils.  Siirli  :iii  cxliact  is  lii'si 
iiiailf  liv  '.'fiiuliiiu'  tlic  entire  ulaiiil  willi  line  sand  in  a  iimrtar  and  tlicn 
cxtractinu  \\illi  a  \\v:\k  (IcciiKiriiial  sdiiilinn  el'  li.v  di'idihiric  ai-id.  TIm 
fxtraci   may  ll'cn  lie  lioilcd.  liltcrcil  tlirini<;li   iiiiislin  and   nearly  neutral 

i/ed,  |irereralily  l.\    iiienns  nl'  s.^liiiin   ; tal(\      IT  ke]i1    in   tliis  acid    reac 

licill,  llic  active  l)rincililc  i>\'  tiie  exlraet  lines  licit  materially  delcrinrale 
willi  lime,  liiit  ]i  il  lie  iieutrali/'d  I'V  coiisidi'ialily  dilii.lcd.  deslriictini 
due  to  .ixidalidii  nceiirs,  ;is  e\  ideiiced  li\  ;i  distinct  liiciu  liiliir  (if  tin- 
soliilimi.  'I'lii'  ai'lixc  principle  nf  such  extracts  has  lieen  isolated  in  .'i 
cr.vstalliiie  Toriii  i 'I'akaiiiini'  and  Aliel'.  Il  iias  lieen  <:i\.'ii  \arioiis  liaiiie- 
''adreiialiii,  siiprarenin,  aclreniii,  (tc ',  Init  tlie  teiidcney  is  di'linilch 
toHards  the  Use  nl'  epincplirine.  ( 'lieiiiicallv .  epinephrine  lias  licen  t'oiind 
to  111-  (irtliodioxyphen\  letliylolmethylamiiie. 


Iln 


no 

/ 
\ 


\ 


■^(1!  nil       til  Micir 


it  will  lie  noted  that  it  is  c-losrly  related  to  tyrosine  I  sec  pay'c  (ill4 1 .  I' 
is  also  closely  related  to  a  iirniip  of  siilistances  lamiiiesi  (U'ciirriiiir  i' 
]iutrid  meat  and  to  which  the  actixc  iniuciiiles  ot'  erirot  lii'hiii<r.  Ii 
contains  an  as>iiimet  ric  carhoii  atom  i  asleiiskecl  in  formula"!,  Mhich 
indicates  that  there  must  he  three  \arieties  of  e|iilieplirille,  dilTeriirj 
from  one  another  in  the  elTect  \vliich  lhe\  produce  on  the  iilaiie  ol 
)tolarized  lisrlit   (i.e.,  a  dextro-  ami  a  levo-rolntory  and  a  racemic  form 

Kpinephrine  can  he  prepared  hy  syiitlietie  means,  the  fiist  |iroduct  oi 
this  synthesis  liciim'  the  rac-emic  salt,  wliii'h  can  then  he  split  h\-  appro 
jiriate  methods  into  di'xtio-  and  levo-  varieties.  The  levo-  variety  ap 
pciirs  to  he  idc'iitical  ill  its  iiliarinacolou'i''  action  with  the  natural  prodiici 
The  dextro-  variety  on  the  other  hand  has  onh  poorl>  developed  ph\sio 
louii'  ;icti\ities  'alio  It  seven  per  cent  that  of  the  l.'vn-  variety^  whih- 
the  racemic  variety  comes  in  lielvveen  the  two  ill  its  action.  A  valuahh 
iissiiy  of  the  aiiioinit  of  epinephrine  in  tissue  extracts  can  he  made  h.^ 
the  method  of  Caiinon,  I-'olin  anil  Denis,'-  in  wliiidi  an  acid  extract  o' 
the  Liland  is  tii  ated  v»  ith  iihospliotnnirslie  aci.l,  and  the  hlue  color  therehv 
developed  eompareil  cnloiiniet  licall  v    with   a  standard   hlue. 

Physiologic  Action 

The  physidhejii-  et'feets  of  the  int  ra\  eiioiis  in.jec'tioii  of  epiiiepiiriiie  an 
mai'kcdlv    excitatorv    and  sliijlitlv    inhihitory   in   nature,     We  will  eonsiih'i 


■Illi;    I  NlHMlil.Vl;    KKOAN^.    <ili    111  111  l>>    (ll.ASl'S 


7:t.'. 


I' 
ll 


tin-  f.rvilalont  iiclloii  firs).  Iinnio.lialcly  iil'tcr  tlio  iiitnivcioiis  injirtioii 
.)f  as  siiiiill  iiii  iiiiKHint  US  0  OOddS  iiiilliirnmis  mci-  kilonriim  of  lioily  wciu'lit, 
11  ilistiiict  ri-.'  ill  iirlctinl  IiIo.hI  i)irssiii«'  iiiiiy  In-  (il.siTvcd.     It  is  (iiiitc  <l<'t- 

iiiitt"   with   (tOOdOS   iiiilli!;riniis   jicr   kiloaiiun,   mid    i-.   ;i< lii|iiiiii.'(l   liy    ii 

>lo\viiiir  nf  tlic  pulse.  This  shiwiny  is  .•aiisnl  l.y  stiiniilalicm  of  the  vatriis 
renter,  as  is  evi.h'iiee.l  hy  tlie  faet  that  if  the  vauiis  iieivs  are  cut,  ..r 
>ilf(ieielit  atropine  adiiiinistere.l  to  paialv/e  llielii.  tile  same  dose  of 
el.inoi)hrine  pi-o.luees  not  a  slouiiii:  |,nt  a  ipiiekeniiii:  of  the  pulse,  and 
eoiiseqiientiy  a  luueli  -iifai.  r  rise  in  Mood  |.r.ssui'e.     The  vaiiiis  a.-tioii 

is  dovelop.Ml  not  lieeailse  of  all  elVeet   of  ei.iii.'phvilK the  va-iis  eeiiter. 

liiil   seeoiidarily  lieeause  of  the  vise  in   lilood   pressure. 

These  pre!iiiiiiiar\  e\periineii(s  indicate  that  the  loens  of  aetimi  ol 
epiiie).hrine.  .so  far  as  the  eiivulatoiy  >\s1eiii  is  ,M,neeriied.  is  iiiainly  on 
the  small  Mood  vessels,  eoiistri.tiin.'  them  and  thus  laiNiii-j  the  peripheral 
resistance.  This  conclusion  can  readily  he  I'ontiriiied  hy  ai>i)lyiuir  the 
el>ilieplirine  .lirectly  to  the  hlood  vessels  of  the  exposed  iiieseiitcry.  ov 
l,y  eiiclosiiii;  a  vas.'ular  oilmii  such  as  the  kidney  in  a  plethysmou'rai)li 
diiriim- the  injection  of  epinephrine,  when  a  ureat  diminution  in  volume, 
acconiiianyiiifr  the  rise  of  arterial  Moo.l  pre^-nrc.  will  he  ohserved.     The 

Misocollstrietinc'  cfi'ect    of  cpilie|)lirille  does  not    liei-niui'  cIe\floped   on    the 

hiru'c  1)1 1  vessels  near  the  heart  on  .•leeount  of  t!.-  deficiency  in  miiscii 

lar  tissue    in    their   walls,      Indt'cd.   these    vessels   may   li< mo   passividy 

dilated  liecause  of  the  inereaseii  lilood  pressure.  The  artiM-ioles  of  dil"- 
terent  ]iar1s  of  the  circulation  are  not  equall,\  sensitive  to  epinephrine; 
those  of  tile  splaiielini<'  area  are  most  sensitive,  whereas  those  of  tho 
ll,. lilt— the  coroiiar.v  \essels  do  iiol  I'cspoiid  at  all  in  most  animals  ("see 
p.i^-e  "J.')?  .  Tile  pu'iiioiiiirv  and  cerelo'al  \c>ssels  lia\i'  a  \ariaMe  reaclivit.v 
lo  epine|)liriiie. 

The  elTect  on  the  vessels  p(  isists  after  complete  destruction,  not  only 
of  the  central  nervous  system,  Imt  also  of  the  vasomotor  nerves;  ei)i- 
iiepliriiie  still  acts,  for  example,  on  vessels  the  nerve  (iliers  of  which 
liaxc  heeii  allowed  to  desieiicrate  iiy  i-uttinu-  them  several  days  liefore  the 
.■piucphriiie  is  applied.  Thi>  would  seem  to  indicale  that  llie  epinephrine 
,M-ts  directly  on  the  iiillseiilar  tissue  in  the  walls  of  the  Mood  vessels, 
liilt  this  does  not  apiicar  to  '>e  the  <'ase,  Wiy  il  has  heeii  found  that  cpi- 
ueplirine  is  incapaMe  of  actiim  on  tissues  wliiidi  are  devoid  id"  sympathetic 
nerve  tiliei's.  and  is  also  inactive  on  those  tissues  in  tile  eiiilu'xo  which  have 
not  vet  received  any  nerve  supply.  In  Iniet',  then,  altlionoh  epine]»hrino 
acts  only  (ui  Mood  vessels  that  are  supidied  hy  tlu'  s.\  iiiitatlietic  nervous 
svstem.  it  is  not  on  the  ner\  >  fillers  liial  liie  cpiiiepliiiiie  nnlolds  its 
actinii.     We  shall  see  iinmedialcly  that   this  ,  onelnsion   's  in  conformity 


•I 


1 


■y   i^r 


m 

!V  i 

1    J    . 

' 

;i,  ' 

15             ;   '     1 

•r 

(•       ! 

'•I  ' 


\  ~iin 


7a(i 


■IMC    I  NIMH  HI  V I      cl|<li\N>.    111!    |)llll,l-s    lil.Wlo' 


\vHh  tlif  n'Mills  III  .iliMMv.iliciiis  iiunlt'  on  sli  iid  hits  ntlur  than  Mic  1)1imm! 
vrssrls. 

Otlnr  imisciilai-  stnictiircs  cxcitcl  liy  cpiiicidii-iiu'  iiif  Jis  follows: 
(1)  the  ililator  uuis.lc  ol'  tlic  i.npil-^.  ispi-.-iaHy  aftor  the  iiorvc  supi.ly  hiis 
l„.,.n  .h'stroyo.l  I'v  .•xtirpatioii  of  tlic  siipi-rior  (■cM-vical  [,'aii^rlioii ;   r2)   Up 

spliiii.-ters  of  the  pylorus  and  of  tli.'  iU>o( at  viilvo;  Ci)   the  must'le  filx-r- 

of  the  spleen,  the  vaiiina,  the  uterus,  the  vas  deferens,  and  the  rotraotoi 
Ileitis.     Refiardini;  tlie  aetioli  on  the  uterus,  however,  it   should  be  notfl 

tiiat    a   different    respon-"    may   1 htained    aeeofdiu<;   to    Avhcthor   the 

uleriis  is  preunant  or  not.  The  plain  muscles  of  the  <M-1)it  and  trlobc  ol 
the  e\e  are  soiiietinies  exeited  hy  suprarenal  extraet,  eausiiig  the  eyes  t" 
protrude,  tin'  pal|ieliral  tissure  to  liecMune  lar^'e  and  tlie  third  eyelid  tn 
he  retraeted,  (dianu'es  whieli  are  very  like  tlio.se  whieli  develop  as  a 
nsiilt   of  frisrbt. 

Inhihilnrii  rlhils  of  i  pim  iihrlin-  on  iiiuscle  are  exhibited  hy  the  follow 
iuL':  n>  the  iiiuselo  of  the  intestine:  ''_'">  the  stoniaeb:  (r!)  the  esopbapus: 
i -\  I   the  i.'all  and  urinary  Maibb'i's. 

Tlie  effeet  <if  epinephrine  in  inliii>itiiiy:  tlie  rbythniie  eontraetions  ol 
,111  isolated  iM.rtioii  of  the  intestine  in  oxy<ieiiated  Rinijer's  solution  is  .i 
\ery  striking  phenoineuoii.  ami  one  whieli,  as  we  sliall  see,  may  be  ver.\ 
suecessfiilly  emi)loyed  for  ileteetiiiir  small  (piantities  of  epinephrine 
Extremely  dilute  solutions  of  epinephrine  increase  the  emitraetions. 

The  elTecIs  of  epiiie)>lirine  on  u^<iii<hiU>r  stnirl nn  ^  arc  tbe  same  as  tbosc 
whi.-li  would  l.e  produced  by  stimulation  of  the  symi)atbetie  nerve  supply 
of  tlie  ..'laud.  Tlius,  the  secretions  of  the  lachrymal  srlaiid,  the  salivary 
udand  (ill  tbe  eatV  the  mucous  friands  of  the  mouth  and  pharynx,  tic 
..riistric  hut  not  tbe  iKiuereatie  -rlr.nds,  .'an  readily  be  shown  to  b- 
ex<'ited. 

From  these  results  as  a  whole,  it   is  evident  that   the  elTect  of  epinepli 
fine  on  muscles  and  -lauds  is  .'xactly  the  same  as  that  which  would  I" 
produced  by  stimulation  of  their  sympathetic  nerve  supply.     This  jiaral 
Iclisiii  of  action  between  e)iiiiei>lirine  and  the  sympathetie  nervous  sy. 
tern  becomes  still  more  evident  when  we  consiiler  certain  of  the  fhdH'l" 
hi    mrliihiillsni   that    follow   administration   of  epineiihriue.     In.iectioii   o' 
e))ine])lirine    excites    i.d.vco!:euolysis    in    tli.'    liver    so    that    hyper^rlycemin 
.mil   -rlycosiiria   become  establishe.l,  results   which   are  also  f.btained  !>> 
stimulatiiii:  the  !,n-eat  splaiudiiiic  nerve.     Kpiiieiihrine  eau.ses  the  elottmii 
time  of  the  blood  discharged  from  the  liver  to  be  very  materially  sliori 
eiied,  ail  effeet  also  prodiieed  by  stimulatiii^r  the  siilaiichiiie  nerve.'" 

As  in  the  case  of  the  blood  vessels,  the  above  results  are  obtained  ev<i' 
after  the  sympathetic  nerves  to  the  part  have  heen  allowed  to  underi;" 
deu'eneratioii,  from  uliicli  it  is  c<,iiclii<Ied  that  tbe  tissues  elabiu'ate  sour 


W   1 


Till     I  \i ;i\i    Mi;.;\\s.  Ill;   hicni^s  (;i  \vi>>  i" 

Mllistillic.  whii-li  ivji.-ls  Willi  rl-.inrl.lirih.'.  'I'liis  Mll.stiilii',.  liiiiv  lie  l.r.)- 
.liiccd  either  ill  tlic  .jilli.-lioii  li.lurni  llic  ihtvc  iiihI  niiisclr  llic  iil\..- 
iii'iiijil   .iiiiicliiiii.     or  |)crli!i|)s   ilir.iiiHliuiit    tlic    priiiu|il;isti!    itx'H".     It    is 

...illnl    the    l-fceplor   Mlh>llUic-e   oT    I.Jiliul.'.v,    illld    is    l.elievc.l    In    iciK-t    linl 

,,lil.\    «ilii   epiiiephriln',   lull    •■,!,,   uitii   Viiri.Mls  .Ini-v      Tlie   I |>lor  siih- 

stlllK-e  S.'fliw  to  ilicrciise.  if  IK'l  ill  Jllliolllit,  ill  least  ill  si'll^it  i\  it  V  lifter 
llie   reilin\;il   (if  t  lie  liel\e  eiiiitliil. 

Kr«..)(.xii.,  wliich  is  :iii  iiinine  ,.litiiine,l  fn.iii  ei-nl  iiii.l  iilso  fr.nii  eer- 

lllin  nf  llie  prodllets  nrllisti-lilie.  lins  nil  Jietioll  CHI   tlie   r. pt..r  snl.stiinee 

whieh  is  illhil.it. .rv  Miel  theiefniv  ;,  lit  il '^ui ,  isl  ie  to  thill  nf  ,.piliepliri,.e. 
The  illllill.M.nislie  ;ieti<.n  of  ,r-oto\ill  Jlffeets  tiie  rNritilt-.r.v  Kilt  llol 
llie  iHlliKltniy  .-leliolis  of  epilieplililie.  ]W  lisilis:  tllis  .llMlL'  WO  :iro  CIl- 
lilileil  1o  slinu-  11, Ml.  iilthoii-h  the  niiiip  effect  of  ..pinei.lirille  oil  tissue  is 
exeitiitory,  ;i  l.'ss  iii;irke,l  inliil.itnry  iiillueiiee  immv  lie  siinilltiilieolisly 
.leveloiH.l.  Til.'  iiihiliitory  effeet  niMy  ,ils,.  s,.nietimes  tie  evoked  l.y 
doses  of  epiliel.hrilie  very  lulleli  sinMller  tlllin  thos<"  used  to  i>rodiiee 
exeitiitory  effeets.  Tiles,,  fiic'ts  iire  well  illllst  int.',!  ill  the  <'ilse  of  tlie 
imiseie  (iiier  of  the  lilood  vess.'ls.     With  iin  onliiiiiry  dose  of  eiMiieplirine 

nmstrietioM  ■urs:  nfter  ervoloxiii  the  s;i„ie  -lose  of  ei.inephrino  eiuises 

diliitiition.  Or  tllis  hitler  result  iiiiiy  idso  be  ol.tiiiiird  liy  iidmiiiister 
in;;  to  ji  iioniiiil  niiiiiiiil  (lUiiiitities  of  epiiiepliriiie  tlmt  are  very 
siiiiiller  thiiii  the  usiiiil  .|iiiiiitity.  The  eoexisteiiee  of  iiiliilutory  imd  ex- 
,-it!i1ory  inllueiiee  is  iils,,  well  iiote.l  ill  the  eiise  of  the  ut.'rus.  In  soiiie 
iiiiiiiiills  the  elTeel  of  ciiiiiephrine  on  tllis  oriiiin  is  to  iUlt,'iiieii1  its  rliylliiiiie 

, ti-iietioiis.  in  others  to  iiiliihit  them.     In  tlie  former  ease,  liowever,  if 

eryf.toxin  is  first  of  all  administenMl.  epinephrine  in  its  usual  dosa<;e  ^^ill 
iuvariiil.iy  produee  an  inhil.itory  etVe.-t.  The  .'r-otoxin  no  doiilit  aets  on 
the  ro.-ep'tor  siil.stanee,  iiml  simihir  effeets  liiive  iilso  l.een  i.ro.lueed  ^\itll 

iipoeodoine. 

Althonji'h  it   is  espe.-iiilly  oli  Jiliiin  iiiusenlar  lilier  liavin-r  a  sympatlietie 

nerve  sui>ply  lliiit  epiiie|.Iiriiie  unfolds  its  aelioii,  yet,  !i< ilin-,'  to  CiUi- 

,„ii,  it  inereiises  the  eoiit liiet iii^'  p..\\cr  of  voluntary  luusele  iind  dimiii- 


•h 


111 

ishc's  the  tilideliey  to  iat  iiriie." 

■|-..r    f.ulli.r   .hl.ils   ..,■    ,Ihm.    ,11....    ,l„'    |.r,;..r.    ..f    I  l.-l.ir-'-    :  ,..M  h.l  IM  .„.■■    -li.-iU    !.. 


'ii 


:•-  I 


3 

5 


t| 


uh,..l 


fl       it- 

IS 


il 


II 


'i 


i;ii  '  it 


■f     • 


i    ' 


■if;!i;''  '1 


i  [ 


^i      '  ! 


pi' ' 

^ 

i     \  \ 


'[  ) 


Ji,x 


(  ll.M'TKi;   l..\.\\ll 

I  liK   ADIiKNAI,  M.ANItS    C.niM 

Variations  in  Physiologic  Activity 

Siiir,.   il    is  clcMl-l.v  i'vliilili>lM'.l   tluit    tlic  iiillviMl   -1:111. Is  ;ir.'   iiidispiMisMl'l' 

f(,  111'.'  iilid  Ihiit  .Nli-ii.-l-.  i>f  III. ■Ill  liM\.'  i\  v.  r\    |.r..ii..i .'.1  i.li\si.,l,.L'i.'  M.'ti.iii. 

it    rciiijiiiis   1 iisi.l.'i'    wlntli.r    111.'    'jhni.U    pr.i.lii.'i'    tliis   iiili'iiiiil    scTf 

ti.ili  «illiili  ill.'  l..i.l.\,  .mil  il"  v..  uli.'tlirr  it  is  rss,lili;il  \,<y  tlir  «  .•ll-lirinu 
III'  till'  iiiiini.'il  ..!■  ir.|iiii..l  .iiil>    un.liT  iTii.-iiii  .•..nilili..iis.     We  must   nls.. 

..iidciivof  ti.   liii.l  ..lit    III \\irh-li    ..I'  tiir   l....lily    run. -li.. IIS  ..I'   III.'   int.'i.-t 

:iniiii:il  tli.'  imnnal  s.  ,-r,'t  ion  .-i.-ts.  Tli.'s.'  |.i'..lilri,K  |i,-iv,'  l.<'.'ii  iillii.-kr.l 
liy  tlii-t'i'  iii('tli..i|s  ..|'  iii\.'sli'^;iti.,ii;  1  l.y  .'..iiipjii'iii'^  till'  I'l.iiicplii'in.' 
.■.".iil.'iit  '.I'  simihiils  |.!rpari'.i  .'\tra.'ts  ,.r  tli.'  I'.'slin-  uliin.l  •■iii'l  "(  "ii.' 
I'.'iiH.v.'d  iil't.'i'  M  p.ric.l  ..r  siip|...s,.,l  iii.'i'i'as.'.l  a.'tivitv;  :  •_•  ■  l.y  .•..lln-tin.- 
Ihi'  lili.o.l  iis  il  li.."s  int..  III.'  vciKi  .'.-iMi  r^'iii  'Ik'  M.lriM.'il  vein  and  i\ 
iiiniiiinsi-  il  I'm-  .'piiH'|>liiiiii'  l..\  pli>  sinln'^ic  tests.  Tlii'sc  .-..nsisl  in  .il.si'fv 
ill..'  tlic  l..'li:i\i..r  .'I'  s.iii.'  tissii,.  tliat  is  s.-iisitiv.'  I.)  111.'  .'i.'tiiin  lit'  I'pincpli 
rini',  siicli  as  tlic  inl.'stin.'  ..i-  iilniis,  aft.-i'  ii|iplyil|.j-  tlii'  I)1>".<1  "V  s.'i'niu 

In  it.  III-  liy  inji'i'tiii-  tin'  M I  ..f  srinm  int  iii\  .'iiniisl>    into  iinntlitr  am 

!iial  and  Innkiiiir  fnr  .pin.  plniiK'  rlVccts:  aii.l  ':!'  I..\  alLiMiinj;  tlic  l.lnn.l 
nt'  tlic  adrenal  vein  I.)  I.e  dis.'liar'.'c.l  iin.iei'  cei-taiii  c.m.litions  tlii'.ni'.'li 
tlic  vena  cava  int..  tli.'  hi.....]  v.'ss.'U  ..l'  tin'  same  animal,  ami  nl.seivin'j 
tlic   elTcct    lil'i.chl.'e.l    ..n    eel'tain    plis  si.il..i.i.-    pt'..eesses    wlii.'li    ill    nne    \\a\ 

.11'   an.itlici'   have   Lii'ii    seiisiti/ed    t.iwanl    tli.'    inlliK'Hei'    ..l'   epineplirine 

Tllis  ant..in,ii'.-tiiin  metli.i.l  lias  le.-.ntl\  I.,  en  use,!  sne.•l■s^|n|ly  l.y  Ste^^ 
,'ii't  and  I{..'J..IV.'"  tlicir  I'avni'itc  sti'ilclnre  nprni  v'lii.'li  I..  ..I.seive  'li. 
epinc]>lllille   elTe.-t    l.eill.^   tile  ilellcl'vatcd    plipil. 

Assaying  the  Epinephrine  Content  of  the  Gland 

AVitli   |-i"4;ii'.l  1.1  llie  tiist   iii.'iiti..ii.'.l  ..I'  llie  m.tli...ls.  litlicr  .'lieiiiieal   ." 
pliysii.l.i'jie    mi'ans    iii,'i>     I.e   eiiipl.iy<'.l    t..    assay    the    streiiL'tli    ..I'   the    es 
irai-ts.     'I'lie  li.st   .'liemi.'al  iiicIIkhI   is  that    nf  raiiion.  Fnlin  and   D'nis, 
the    iirini'ipli'    ..f    -  hi.-li    has    alna.ly    lieeii    .Icser;'  ed.      'I'he    jiliv  siol.iL'i.' 
iiictlmd  yicMin'j  m..st   satisla.'t.ny  ri'siilts  is  1li;il   i.t'  HUii.tt,''  uliieli  e.iii 

sists   in    injeetiim-   a    p.nti I'    tin'   extract    intravenously    intn   animals 

t'roiu  which  the  intlneii.'.'  ..i  tiie  nei'\.'  centeis  nil  tin'  heart  and  I.!....-; 
vessels   has    I. ecu    i'ei.i..\e.l    li>-   .leca pitat ii.n.      The    risi'    in    ai'lcrial    IiI.k.'I 

73S 


iiii     Mii:i  \  \i    1.1  \sii^ 


:!'• 


].irssUff  |.fi"llliT(!  li>  till'  ih.irrliiMi  is  lliin  .-i  \r\-\  I'iiii  liMiisii  I'l'  »f  III'' 
.iiiiniliit  ul'  f|iilli'|>liiinr  c.(Hil;rnir,|  in  il.  Il  li.i-  l»i'li  -ti""  li  itiiit  llic  !•'■ 
Mills. ilitiiiiifil  l..\  llii'i'lii-niii-iil  iiii'llic"!  ii-rii'  v.tv  .InMly  willi  tlms.'  ditaiiinl 
liv  111.'  |iliysi.,|..._'i,..  lull  il  sIm.iiI.I  I'.'  i.-Miiirl«'.l  ili.ii  ■'.  i>  .lilli.'iill  in  s..,.  li,,u-  ili.- 
i.liysi.ili.iii.-  liii'llii"!  .•..niij  \«-  ;i.-<-i;i;ili.  in  all  .■asis,  siii.'c  il  Ikk  ImcU 
sIh.wii  llial  willi  'Ji-.al  .|ih;ii..n  ..l'  .  imnpli  i  in.'  a  iv\.msi,|  .  IV.m'I  a  \asu- 
.lilalali.iii      iiiav     \>r    ..hlaiii.'.l.      .\nii.i|,U    in    assa\     tli.>    stn-n.'tli    nl'    an 

.■|iilic|.liriiii'   si.lulinnj.\     iiiMsli'jaliiii;    llu-   .■IT.'.'N    wlii.-li    il    I'I'mIi «,   nii 

.illicr  pifpaialiHiis.  sii.-li  a>  is,,lal..l  \<«>]<^  "\  nit.Ntin.-  ..r  ul.Tils,  nr  iIm' 
.iiili'lcal.'il  .'.\.'liall  .il'  111.'  irnu.  niusl  \<\n\,'  uiiMi.-.'.-.srii|.  >in.'c  tli.  cir.'cts 
arc  liy  im>  iin'aiis  .{.■|>.'iiil.'iil  iMi  llif  .-niiriiil  rat  i'Mi  <,\  .■|.iii.>|iliriiic  in  tin' 
.Alract.  WliiMi  sii.'li  |>ri'|.arali..ii-  aiv  iis..l  h.r  ,|nanlital  iv,'  |iur|iiis.-s. 
llic  sir.'iiulli  ni'  llic  .-strai'l  iiiiisl  Ih'  JilI'^.m!  l.y  IliMliiii;'  lli.'  .aLmiI  I" 
wlii.-li  il  .'an  I...  .liliil.'.l  ami  siiil   r.iiiaiii  a.-li\.'. 

(^Uitf    a|ial't     rinlll    till'     I'nICUnillU     |Mis>il,|..    -,,  ,11  li'i's    nl     rfVi^V,     it     IMIIst     li-' 

i,iii..|iili.'if<l  llial  llic  nsillts  iiiir.'l\   ;ji\c  ils  an  i'l.'a  nf  Imw   liiu.-h  cpiiicpli- 

:iHc  iiiax    lia\c  lii'cii  lain. '.I   in   ll;<'  'jlaii.l   at    lli.'   time   nl    its  .A.-isLni. 

Tlii'N  i-an  nut  ti'll  lis  Imu  iiiin-li  .■|iiiiriiliiani'  llic  ^laiul  was  s,.-r.'l  iii^r.  I'lim- 
111  .  \.-isi.in  as  iiiii.-li  .if  ti'is  ImnicMic  iiiiLi'il  lia\  .■  Iic.'ii  lin.li'i-'j.iiii'-;  a  prm-.'ss 
nf  iiianiirac-luri'  in  llic  lilaml  as  was  luinu  iliscliai-'_'.'il    ri-mii   it.  sn  llial   tli.- 

.issav.'il  aiimmil  \v..iilii  ivpivs,.|il  iii.svIn   tli.'  I.alam I'  pn"lili'linn  ami  Inss 

,,,■  lioni: Ii>    llic  '-'land.        \V,'  mi'jlit   .piil.'  well   lin.l  that   tli.'  aiii.ninl   "I' 

..pincpluiiii"    in    llii'   .a. 'is.', I    -laiul    was    mniiial    iiii.l.r   .•nii.!ili.>iis   wticri- 

tli.Tc  lia.l  lie. '11  an  c\i'cssi\..  .Iis,.liaii;.    ..f  it   int..  tli.'  M 1:  tliat   is  1i.  say, 

Inss  and  pi-.Mlii.'ti.in  mii;lit  lia\.-  la-.ii  .■.|ii.'il.  Wli.'iv,  Imwcvcr.  a  inarkcl 
.Icli.-icncy  is  rmin.!  I..  c\isl,  il  pml.alily  i'cli.-at.s  lliat  exhaust inii  .it'  the 
]inwcr  .il'  pi-n.liieinu   cpii i.' pli lin.'   '^as  lakiiii;    pla.'c. 

The  Epinephrine  Content  of  the  Blood,     'llic  s, n.j  imilm.!,  in  wlii.ti 

I, I,,.,. I  |-r,im  animal  is  tist...|  i..r  its  cpin.pliiiii.'  ciV.'.'l   liy  intra' .■imils 

iiij...'ti.)ii  int..  aiintli.'V  animal  nr  \,\  appl.\  inu  it  tn  s.mi.'  isnialc.l  pr.'paia 
tinii  nil  wl'icli  cpiii.'plii-iiic  a. -Is.  has  \i'l.|.'.l  impnilani  csiilts.  Sin--e 
.,.■111111  contains  all  the  cpincphi'iii.'  .il  li|nn,|,  it  ..an  !..■  c..nvcniciilly  usc.l 
inr  111.'  tests  'Stewart  ami  IJ.i^nir  .  Tim  is,.lat.'.|  pliysinl..i.'ic  l>i'ci>ara- 
ti.ms  that  hav..  Im.^ii  uscj  i,  ii'stiiie  \\„-  ..pin.plii-in..  in  ih.'  aniiii,-il  llui.l- 
ai-.'  as  I'.ilhiw  s: 

I.    .1    s-ilnl'lfl    uf    II,,     s„n,ll    nihslnl,     ..|'    a     fahhil.    s|is|  „.|i,l,.,l    ill    nxy.Jl'll- 

ai.'.l  |,.M-ki'"s  s.iliitinii  at  lin-ly  t.iiipeialii!'.'. 

•_>.   .1  si'iiiKiit  nf  t!u    ,if<  riis  ,,(  ■.{  n.iiipi'i"jnaii1  rahhil   siinilai-l\    pri'|iavcii. 

Til.'  apparatus  iisi.,|  f.ir  .ihs..r\  in.^  tli.'  .•.ml  raclhuis  ..l'  citlicf  i>rciiara- 
li.m  cniisis's  111'  a  small  ulass  .•liamhcr  riirnisli...!  h.'h.w  with  a  li.mk  in 
which  ..lie  .'ml  ..r  Ilic  s.-uii'iil   is  attach..,|.  Hi..  ntli..|.  ..n.!  l..-ilii;  .■mnu-.-t.-l 


in  i 


•        •     i 

1*' 


!      }i^ 


740 


III     IMHM  ItlNi:   ORIUNS.   UK    Ml  (TI.ISS   (lIANHS 


to  a  niuscli-  Irvcr,  mi  tliat  llic  rcuMliir  rli.Mliinic  r(.iilia<-ti..ns  can  I"'  ivuis 
toroil  on  n  drnni  (h'itf.  100). 

Kltiin-Iihriiu'  inliil.its  tin Mtra.-ti<ms  «(  tlio  intostino  lint  stimulalo-i 

llioso  of  llif  iitiius  of  most  aiiiinals,  tin-  iiitcstinc  iii-cpaiatioii  \mu>i  \h<- 
nioro  sonsitivo  (Kin.  1!M  i.  Imlti'il.  it  is  said  llial  tlii"  iiiliiliition  in  tlii- 
caso  inny  bo  olitaiiicd  willi  a  snliiliini  coiitaiiiini.'  1  pail  of  cpiiMpliiinc  in 
'20,n()(),fUK)  of  solution.  In  iisiiit;  tliis  niotliod,  liowrvcr,  !;n'at  caro  and 
juduiupnt  ninst  l>i'  exorcised  in  dniwiiiii  conclusions,  liecause  other  siili 
stances  present   in  the  lilood  are  lialile  to  affect   tlie  contractions;  thus, 


Air,  renf 


l-'ie     191J       ;\rraiiBi-iiitiil    i.f    aM..1i.ilns  f..r    rcinriiinif    .ni,tr:uli..iis    ot    a    ulrri  i.'    -Ini.,    niti -Ir   r 

slrip    or  ring    <tc.     Thi-   nirlal   w^il.r  lath  is  nia<lf  of  a  ili.ap   nutal    wal.r  pail   «illl   a   li.almt!   m, 

s..l,lVre(l  Ihr.u.Kli  the  si.lf  .it  the   l...ti,„M.  A   shnrt  im-tal   tvil.e  is  soMcn-.l  into  a   ImkIi   .M'.oniu   ' 

the    bottom   to    recrive   a   Krforate.l   crk  for   coniiciting    with    the    llarvat.l    iinisrlc-v.arm.r    niM.i, 
(From  Jackson.) 

certain  siibstanees  in  blood  seniiu  which  liavc  been  i)rodnced  l>y  the  lo' 
of  lilood  clottini:  may  cause  auvnncnlatimi  of  the  beat  in  both  the  iiili^ 
tinal  and  the  uterine  preparations.  A  certain  amount  of  eitinephrine  m 
Locke's  solution  is  con.sc<|nently  more  likely  to  cause  inhibition  of  tin- 
intestine  than  a  similar  amount  aihlcd  to  blood  sennii,  because  in  the  lat 
ter  case  the  jircssor  substance  will  iieulrali/.e  the  depressor  effect  of  tin 
epinephrine.  On  the  tilcrine  jneparation.  both  tlie  blood  serum  and  He 
epinephrine  have  pressor  effects.  As  has  been  pointed  out  by  0.  N 
Stewart/*  if  both  prepaiations  are  eiii])loytM  for  testing  a  solution  suj) 


M 


5\1 


Tin;    ADKKNM-   (II.ANIM* 


741 


posfd  to  ronlaii.  i-pin.-pliriiH-,  liltlo  clmiirc  of  niof  is  likely  to  1)C  i»- 
ciirictl;  tliat  in,  if  tiio  soliilioii  iiioducos  iiiliiltiti«in  "f  Hit'  intestine  nloim 
with  aiii.'iiiciitiiti'>ii  of  llic  ut.'ius,  it  must  e.intain  epinepiiriiio. 

3.  The  frrxh  nimlid  arlfrii  of  Un   simi,.     A  riii^'  eiit  fn.m  Ilic  artery 
is  siisp.MHle.l   ill   oxy^'enafe.!   I.o.-iie's   s..liiti<.ii   aii.l   attiicheil   I'elow   to   a 


f  ? 


|-,„     I'.l      Tracins   sh...in«   Ihr   eflcct   of   .rinei.hrinc   on   the   intestinal    "'''"''.'r';'   '."','   ''','l,,','!'' 
,.t,n"l   l'l..';.l   ,'r,-.Kr,'.      iTlu-   pr.  Inn. nary    a.l.lili-n   ..f   Larnnn    lo   the    nutv.t.vc   fluul    nw>    U-   'I1-" 
::  .r.li.l.l       (  I'r.iiM    .laik-^.n.  1 

Muall  iHM.k  an,l  abnx.'  In  u  li^iit  nius.'l,.  lover,  l.y  xvliidi  llie  <M.iitraetion 
of  the  iiiiisclc  filifis  ciui  he  ohsiTve.l.  Kiiiiiepiiriiie  eaiises  liu'  muscle  to 
contract,  but  the  test  is  not  so  sensitive  as  the  forejjoing,  especially  in 
the  presence  of  blooil  serum,  because  the  pressor  substances  therein  oon- 
Ijiinetl  also  cause  contraction.  I'.loo-l  l>lasiim  docs  not  contain  the  pres- 
sor substance,  so  that  oxahilcd  I'lasiiia  sjioul.l  be  used  in  place  of  scrum 


il  i  I 


7I'J 


I  III      I  •.hill  HIS  i     mil,  \S-,    nil    111   1    I  II  -^    1.1    \  Ml' 


ill    :i|i|ih  ih'.;    llii'    ti>l        'I'll    Mini'.ix     I  lie    xnisil  Imih'^-.    hI    iIii-    iini-rlr.    lln 
llllcrv     lillL'    nImiuM    111-    >lli!llll\     -Inirllril    li\      InilililliJ    lllr    \i\i\- 

I.     Ill'     liliii'il    ri.>.'>    'il    .'    '""/        'I'lic.    hii'tlliul    i|r|ii'lliU    nil    lllr    ^illlir    |inil 


'if  •! 


i'- 


-  S 


lili 


■i^'J 


i-ijilr   JIN    ill    tli:it     ill^l    ili'«i-riliril.       'rilr    Mllill    -II  j  ij  li  iti  1    I Illllill    rjii  In'llll  ri  1 1' 

i,    ailili'il     ti,    Lim-Im-'-    -.ilulimi.     whii'l,     i>    iM'Miiu  lill'-    iiriii'j    |iiTlllsril     limlil 
i-,;,llil     |i|-iN-.illT    thliiuvli     llir    liloiiil     \(--Ni'U    iiiiil     llir     liilr     111'    iiUtl!ii" 


nil      VIilll  N  M     '■!   ^"^1" 


743 


n..tr.l  .l-'i'.'     !!•■-'         II'  III.'  Illll.l   JMl.lr.l   t..  II,.'   Ihltnuill..'   llni-l  -•-'iiliiii's  '-l-i- 

„..|.!,rin-.ll lltlinu    ulll   lMrn,Mr.|llMinislM..|.      Tlil>  is  11   x.M    si.  ( isllirl  nr> 

„H.|lH»t.     Jll.uuull     i-     i>    > "l''"     '"""''I    '"    ■"■"'"•    ""''■"'    ''"■'■"'    "'"-^    '"■'■ 

,„ ,,'l,i   .    l„,.au>.'    nl'    Ihr    .lirti.M,ll>    ..I    .i.nin..'    tl,..    i,r,-.-..nr>    .mhimiI..' 

Iiitn  111.'    x.vmU     iiniln   iilhl   al"|.iinii'iil    \''in  •- 

,-,,    ■II,.    ,,„,„!  ,.l    th.    <,nn!,,it.,l  <■!•    "'    ''■'     ''■■■''.      KMlv I>    s'llMll    tni.'ps 

,,f    ..|,il|r|.|,nMr    111.-    ..l.-riv.',!    In    .•;n|..-    .1    .hlatMlM.lt. 

,;    •//,,   ,l..u,r.,i.,l  .-'-v     Tl,..  iIuhI  t„l.,   I.M..I  u  |,l.....l  in  tl .jniM-- 

,lv;,l   VI.-  ..I    i llnal    fi..ln    ulu-l,    ll,:     >u|..ii..r  .■.■'vi--Ml    •-'an..-liHi,   nf   til- 

,.,„.,■...,.„„.!, n.  .i.l,-  liH.  l.-.n   I.- v-i  -m,r  ,i,,  v  ,  |..vv  i..i|.h  ,      I  n-l-r  ml    | 

, |„i,„„,   M  .■,.,imH,m,u.  i,  lu.M.n,    in  tL.   ,lni.|,  .lihitiition  ..ft),.,  l.iipil 

,„.,-iirs     i;..iii  .  r  iii.  |.i..-..iiii-  i.a.'ii.'Ms  uf  .,^v.'  t..  M.-H/.T. 

I,    ,1 1.1    l„.   ..,M|.lia.i/r.l    lii.-.l,    alllinu.J,    .a.-li    ..f   tlir^r    niH Is    i.    HI 

ilsrir    V.r>     s.MIMtiv.'     t..l'    til.'     .|.t.'.'ti..M     ..I     ,l.|IH|.|.lill.'     UltlH.Ul     l.nilU:    111 

„nv>   s,M.-iti..,    v..t    tl,..    i.>nlt    >1 .1    ti"l    1 nsi.i.MT.l    iMsivr    lllil.  >s 

.,,.,•„,•„„    Hr,..-ts   liav-   iH.ii   -a-iiivl    I'V   Mt    Last    tu,.   ni.-tl,...K   tliat    ar-   ;is 
far  as    |...-,sil,l..    in.lr|>"li.l<iit    ..f  ra.-li    ..tli.r 

V,  a it.'...ii-  ..r   ii,^.str.ati..ns  l.v  tli.M.    ih..I,...1-.   It    l^s   U,n    fniiii.l 

,|,al     ulirn  1.1 1    f. hr  M.lnmil   ^''^n   ,s  ....ll.Ttr.l  in  a   1 K-'t   "I'  vnni 

>    al.'.v.'    :iihI    l"'i"U     tlif    riiliilllf.-    nl     111'' 


a\a    iiiailr    li\    a|.|.l>  mi;    I'laiiij.s 


I.,.   r.\.al.'.l.   tlic   nitc  "I 


a.ll'4'liiil    v'iiis,   tin'   l>r.'s.i r  c|,ii,.-|.liriin'   I'.-iii 

.,.,.vrti..li    l..'iiM-'    fn.lii   tMHlii:;   t,.   (ttHtl    IM'..    \«-r    'ulnuniM.    '<(   IhhIv   «.'lU'lit 

,,„,  ,„i,H.t,'     Sl.uart    aii.i    l{..-.ir         Th-  al.s..lul,.  at mt    ..l'  -l-H-I'ltnin' 

lil,..rat.-l    fn.iii    tin.   .Jan.!   .■an    !..•    in.asurr.l    ..nly    l.>    titul.n..'   tl,,. .'H- 

„.ati„.i  in  tl,.-  a.ltvnal  s,„,  M I  mi,.I  tl,.-  lat,-  .,f  1.1 ll!'.".     Tins  annnmt 

is  ai.l.,-..Nn„.,t.-lv    .-'-^'''t,  s„  thai    tl,r  ,-„„-,.„ti-ati.,ii   in  tlu"  I.1."m1  ^^ltt'•l' 

,.,.ll,.Hs   i„   tin-  .'-ava    , k.  t    vaii.s   invis-lv     uitl.   tl,-    ,-:it-   of   M Hl-W- 

In   .,s|.l,^-.,.    tl,.-    1.1 M.-"    is   .!.■.-,■  as.. I    >,.   It, at    ll„-   .-.., nt,-at„.ti   nt    -pi 

„  ,,,in-  „„-,-as..s.  iMiI  tl,,-,-  is  „.,  -Inn,.,-  in  tl„-  nl,s,.|i,t-  a.i.n.itit.  N-i- 
,1,,,,.    a„-stl,.-s,a    inn-    tianina    alV-.-ts    tl„.    ai„..iint.      Tl,-    -nl„-t,t  rat  i,.,,    |s 

lik,.|v  t,.  IIS-  lat-  in  an  -s|,-i  inn-n.    l.-.nis-  of  tl,,-  si,.,  iii.^  „f  1.1 Itl-.w 

\,lmwil  ii-tivitv  l„a^.  linn.-v.-i-.  1-  -x-it-,1  !•>  'nassa^,-  of  tl,-  t^liiiwl.  "f 
l,v  stiniiilalinn   of  its  i,-n.-  supfl^    tl„-,.„..l,    lb-   un-iH    splan-liiii-   i,-n-^ 

Tl„.  pi--s.„,-,.  of  .pin.  Phfiii-  in  M 1  .-..H-t-l  'lii Hy  from  tli-  .,lf-nMl 

^,.i„s  ,l.M-s  „.,l    instilv    „s  in   -,.n.-lu.lii,.^   tl,al.   v.l,-n    iiiiNnl    "  itli   tl,.-    f- 

,„,,-|ii.l.-i-  of  tl,-  1.1 1  in  tl,-  l....l>.  tl,,-,-.-  u..nM  I'-  .-1  snir,.-i-iit  -oi,-.'iiti-a- 

,„,„  „,-  ,|,i.  snl.sta,,.-  t.,  ,l,-v.-l..p  :in>  ..f  its  a-liviti,-s.  It  l,as  tli-n-tor, 
1,,,,,,  „.-,-ssai-v  t..  .1,-Ms.-  ii,.-tl,...ls  l.v  ul,i-li  tins  p.,ssilnlil.v  .-oiiM  1- 
tcstcd. 

The  Autoinjection  Method.    s,i,-l,  a  ip-i',o.I  vmis  (ii-si  of  nil  sikv-ss- 

f„llv  iis.-,|  l,v  ,\s|,..,-.  ul,..  .-inpiov-i  :,o  :'""-•'  ^  -I't  ^hu-h  all  tin-  a'"  o"' 
,„,,lvis..-i-a'l,a,l  1 1,  i-.-iiiov,-.l.     ft,  siinmlat,    ,,  ..f  th-  ..r.-at   splan-lini- 


M  il 


1  f 


^l\v'  ^'i 


m  . 


lilt 


■;li 


I  i  •! 


Ui    I 


744 


iiii:  iMiiii KiNr:  (iiii;\\>,  oi!  h:  i  ■ii.i,>^  (;i,am>s 


iiiTM'  :i  ii>i'  ill  .•iilcii;il  Miiiicl  |iiiN^in  T  nrciiirc'cl  pmviiliMl  tlic  iiclii'ii:il 
\cilis  wen-  oprii.  Iillt  lint  s(i  il'  llic  ;iilrcii;il  xciiis  were  cliiniprd.  \\\  li- 
iiioviiij.'  tlir  \  i~^cfi;i.  llu'  ilTti't   iif  ^i>l;iiii'linic  stiiiiiilatioii  mi  llio  mImIoih 

iii;il   111 1   M'SM'ls  llii'iiischrs   is  I'liiiiiii.-itcil.   lunl   .'iiiy  ciiiisl rict imi   wliic'li 

cJCClirs   ill   tlir   lilno.l  vessels  ii|'  tile   list    cil'  tllc   \>m\\   IIIMsI   dhviollslv   lie  illle 

tl)    till'    ill'linll     Ml'    f|)illC|lll|-ilM'. 

'I'lir  iiKPst  salisl'iii-tiiiy  of  llirsc  iiiitlinils  is  that  iimrt'  iccciitly  eiiipInN  i.l 
hy  StcHait,  l.'ducilT  and  CiliMin,'  '  wliicli  ciPiisists  in  dliscrviiiK  tlic  In- 
!ia\i(ir  nf  tlie  |niiiil  on  tlic  side  iVnni  wiiii-li  the  siiiicrior  ■■ci'vical  u'aiij;li"ii 
lias  liccn  ic'iiin\cd  aliiMit  (piic  wirk  prcv  imisly.  (  M'  rniirsc  tlu  l)li"id  pri's 
sure  oll'c  rl   Is  alsci  iilisi'i-\  III. 

AnmiiL;   ilif  must    iiii|"niaiil    rcsiills  Ncciiicd   li>    liiis  Mhllmd   it    may   I" 

•iii'iiti cl  t|i,-it  dilatali'in  (.|'  ili.>  |in|>il  nciMiis  on  st  iiinilat  ioii  of  tlic  iricai 

s|ilanclinir  n"r\r,  |irn\i  i-d  tlir  \rna  cava  and  adri'iial  M'iii  aii'  unolivt  rilc-tid 
so  that  till'  lijond  t'rniii  till'  adii'iial  ^daiids  i-aii  i;i't  to  tlit'  lii'ad.  If  the  \i'Im 
ra\a  is  claiiiln'il  alnl  the  s|ilali(dinii'  iiitm'  sI  iii;iilatc(l,  tiicif  is  no  pupil 
lary  dilatation,  Inil  it  iiiiiMC(lialt'ly  orniis  allcr  the  damp  is  i-cirio\cd 
I';piiii'l)lii'inc  i-oni  iiiM.s  to  lie  disrliai-iicd  lor  ii  cniisiiliM-al'lc  pnioil  ol'  tiim 
at'lcr  stiiiiiilal  iiiu'  llir  sphiiiclinic  nc-r\  e.  lull  llif  imiindiati'  iiicirasi'  w  liirl 
I'.illows  till'  application  ol'  the  ^tiirinliis  dors  not  la'^l  loiiir,  so  tiial  liini. 
accretion  .'.Hi  Pc  nlitained  Py  inlcriiiitleiit  than  1>\  i-oiit  inuous  stimnla 
tinii.  Il  does  not  sei'iii  to  lie  po^sihlc  to  e\liai|s|  the  adi'elial  L'laiid  o('  il- 
suppl\     of    ,ieti\e    inatelial     liy    si  imlllat  illii'    the    splallehnie       ;|     fact     W  lliel 

would     seem     1.1     thlow      iNiijcralilc    dollht     on     tile     reliallilily    of    till II 

elusions  ai-li\ed  at  li\  the  Use  of  those  iiielhoijs  in  wliicli  exiiacts  of  tin 
'jland   ale  :  n-;i\..|      -.,•.•  |.;|.jc  7:'.''    .' 

.Maii>    iiitei-est  ini;'  facK  I'oni'iTiiiiiL:  the  iialiii'c  of  tiie  iiinei\ation  ol'  tie 

ulaiid   have   hecii   si', Mired   \>\    o ■  oilier  of  tlic  alioxc   methods.      After 

section  ol  the  sxmpathelii.  ihain  and  the  v:reat  ■lilaiiehnic  iier\i's  on  holh 
sidi'S  'in  ihe  tl,.na\  ,  no  epinephrine  is  secreted  into  the  Mood  of  lie 
adrenal  vein,  and  whin  one  'jland  is  extirpaleil  aiicj  the  iier\c  connee 
tiniis   of   the    other   entirely    cut.    the   epiiicplirine   content    of    tlic    adrenal 

vein   liloocj   sinks  to   not    i -c   than    1    liM'il  of  the   inn-nial   amount,     'i'lo' 

animals  siir\i\e  this  hillei-  operation  ami  liehavi'  in  a  perfectly  nmiiia' 
fashion,  inilicatinir  I  ha  I  the  internal  secretion  of  the  adl'i'iials  can  imt  ha\  e 
the  iili\  sioio',:ic  siirniticance  so  often  asiM-ilieil   lo   it. 

The     s|il,|||i-|inic     lilicls     iMiliei'lneil     ill     ihr     sc-letiol!     of    epi  I  le  |  ill  ri  ne    seem 

li nie   fioiii  a   nii\e  i-eiiti  r  situated  relali\el\    lo\,    down  in   tlic  spina' 

eolil.       Seetioll    of    llle    cord    at     llle    IcXcl    of    the    last    cervical    seU'nlCIlt    lloes 

not  atVcct  'he  spoilt  a  molls  secret  ion.  In  it  this  disappears  w  hen  the  sect  ion 
is  niailc  lielow    the  third  thoracic  serine  it.       (Stewart   and   KouolV.I 


■I.  \.  l.-j  Liiinl    ..1 


ll....-:'l.     .1..II 


'       I         :    ! 


Till:    AKHIN  M.    (il.AM)S 


74) 


III  cuMn.Ttioii  with  tluM'  ol.scrvatic.ns  il  is  of  int.'ivst  to  note  tliat  tliif- 
iiii,'  stiimiliitioii  of  Die  spliiiwlini.'  iifivc  in  a  iioniiiil  iiiiiina!,  tli«>  (■"iisc- 
,,il,.|it  rJM-  ill  IiIchmI  prcssiiiv  slmus  t«o  peaks  i  see  l-'i^'.  -JO.  pa-re  l:!Ti.  Tiie 
first  is  HO  ilniil.t  line  to  diieel  si iiiiulatioii  of  tlie  splaiieliiiie  vasocoustrie- 
lors,  and  '.■•  sei  c.ikI  to  tlie  niilpoiinnii  of  epinepliiine  into  |lie  1,1, .od,  tlie 
i,isti''-.i>:..:'  S.r  iliis  eoiichisioii  lieiii'j  that  the  latter  risi'  Tails  to  appear 
al'le     !  riMo\  al   ■•!   tin    ..  Ireiia!  u'lallds. 

•|',  .;,  ,  i|..  i.-ilt  .IS  a  whole,  it  is  indeed  doulitflll  w  lieth.T  Ull.ler  llol'- 
iiial  .■  ii.'iti'ei-  a  M.jieient  am. .lint  o|'  epinrphiin.'  is  diseliaru;ed  into  the 
Mood  of  the  ve.ia  .ava  t(^  alVeet  api'feeiahl.v  the  tone  of  the  hloo.l  vess.'ls. 

,.ind  this  <•( hisioii  seisms  all  the  more  .instilled  heeaiise  .-f  the  fact   that 

small  .piantities  ..f  epinephrine  h-.wv  a  .lilatinu  rather  than  a  e..iistrielini;' 

iiillneii,-,..  .-It  l,.asi  ,,i riaiii  vesxls  I  ll.irtmair  '  i.    It  may  \y.-.  howex.M'.  that 

Ihe  iiiainlenaiie.-  of  vasenlar  tone  under  certain  conditions  is  irreatly  as- 
sisted l>y  Ihe  presence  of  epinejihrine  in  Ihe  Mood.  Siinilarlx  the  syinpa- 
thetie  eonlrol  of  other  functions  may  he  facilitated  l>y  the  presence  of 
small  amounts,  it  has  Keen  found,  fur  example,  that,  aithoiiudi  stiniula- 
linii  of  the  c'cliac  j.lcxns  causes  tlie  -l\  co-en  stoivd  in  the  liver  to  ho  eon- 
verled  into  su-ar.  this  result  is  not  as  a  rule  olitained  on  stiinulatini.' 
l.h'xns  shoilly  after  removal  •'(  the  adrenal  •riands.  The  in-esence 
,,1  epinephrine  in  the  Mood  u,,nl.l.  thercl'.u'e.  veem  tol...  necessary  to  l)riiijr 
Mhoiit  fnnclioual  a.'tlvit\  of  the  sympathetic  nerve  en.lin-s  eoncenuHl  in 
till'   ^d>  ciejeiiii!;  1  il-   pi ss    :  vee   pa-e  I,:',,     . 

Adrenalemia.  In  the  liuht  of  these  researches  ii  is  important  to  point 
unt  that  a  ureal  part  of  the  uork  done  l,y  clinical  ol.servers  i.urportinsr  to 
show  that  in  such  conditions  as  nephritis  and  arteriosclerosis  there  is  an 
increase  of  epinei.hrine  in  111.'  Mood,  has  hecn  foiiinl  l.y  Stewart  and 
others,  nsin-  controllcl  methods,  to  he  entirely  nupro\.n.''  Some  iiiyes- 
ti-atofs,  however,  still  In.ld  that  temporary  conditions,  sndi  as  transient 
ris.'s  of  .'irtcrial  Mood  pressure  or  temporarx  ulycsuria.  max  sometiinos  l>e 
dne  to  inci-casi'd  adrenal  <liscliarue  into  the  Mood. 

Kphinephrine  has  hecli  tholl-hl  to  he  a  suhstanee  which  is  sccrete.l  into 
IheMooii  in  supernormal  amount  when  certain  eiiierireneies  arise,  the  most 
i,„,,„rtant  of  these  hein-  fright,  or  srnne  other  extreme  emotion.  This 
helief  has  arisen  partly  from  the  similarity  l.elwcen  the  jieiieral  hehavior 
of  an  animal  follow  in-r  the  inl  ia\  enoiis  injeelion  of  epinephrine  and  dnr- 

iiisr  stales  of  extreme  excitement.      Dilatati f  the  pupils,  hrisllinn  of 

the  hair,  salivation,  rise  in  arliuial  Mood  pressure,  inhiliition  of  the  intes- 
tinal movements,  protrnsioii  of  ihe  eyehalls  are  all  symptoms  of  fear  .just 
as  they  ar>'  of  e|)inepliiine  injection.     Impressed  hy  these  resenililances 

i'^nnun-  undertook  an  ,■•.!.■'. l-l  re-;!-  !■  '■'  fsl  the  liypothesis  that  the 
ivaeti •  an  animal  to  fear  and  other  emolional  slates  is  .lepemlenl   on 


in 


\  i    M; 


S: 


Wi 

.    >i 

',U   f  13 


'    1 

in- 


l:M|-JiiiN' 


.* 


1 

t!      i 


,  '-^  :'      I 


Tlii 


rill      I   SIMM  UIVl;    (IH(iANS.    Uli    |)ICI'I,I»-    (il.ANIi; 


li\  ]iiCMTni  inn  111'  i|>ilicplifilic  ilitn  I  lie  MhihI.  TIh'  roll  Its  scfliicil  1()  <'i)ii 
liriii  llii'  li>  |H.ihisi^.  Ill  till'  liist  phici'.  it  was  fmiiiil  tluit.  wIhtcjis  tin-  IiIihhI 
ili-awn  tViiiii  tlir  \iiia  iM\a  iipiiiiNiif  ilic  intry  nl'  llir  ailfciial  \riiis  i  li> 
|iassiiiL:  a  c'at  liclir  up  lln'  I'niKiral  \  riii  till  its  iVcc  cik!  la\    at  this  level  i   in 

; nnal    male  eat  iliij   imt    <:i\e  e\iilen if  tlie   preseii >l'  epiiieplirine 

w  lien  testeil  liv  means  n|'  tile  intestinal  seirmeiit  iiiellid'  it  did  so  in  a 
i-at  lliat  had  ]Me\  inii^lv  lieeii  iVi'/litened  liy  allin\inir  a  diej;  U<  liark  at  il 
Siieli  lesults  \\t\i'  lint  iilitained  al'ler  leiimval  nl'  tlie  adrenal  Lilalid,  m-  ii 
a  t'eiiiale  cmt,  wliieh  is  iisiiall\  indirrei-eiil  tn  siieli  a  meilmcl  nf  friirliti'iiinu 
( 'aiinnn  aisn  tlimiLilit  tliat  maiiv  n I'  tlie  ntliei-  adaplaliniis  w  lii<-li  take  plaee 
ill  an  animal    in   lliis  enmlitinn  eniiM   lie  att  lilnited  tn  the   pieseiiee  nj'  a? 

exeess   nl'   epinephrine    in    the    M I        Tile    thri'e   lllnst    illipiirlaill    ol'   tliesi 

are:  '  I  inei-eased  discharge  111'  s  11  liar  I'inm  the  li\  cr  illtn  the  lilnnd  ;  l  "J  I  in 
ereased  el'tieiem-y  (I I'  museiilar  eniit  raetinii ;  in!  diiiiiiiislied  (dnitintr  tiniii  ni 

the  111 I      all  nl'  w  liiidi  are  adapt  at  inns  eiiahliiii;'  the  animal  either  In  enn 

ipler  the  sniir t'  the   Tear  nr  tn  he   in   a  hetli'l-  ]insilinn    tn   reenver   fnim 

aii\  lindily  injtirv  in\nl\in'_;  a  Inss  nl'  hlnnd  shniild  he  sutTer  lindily  dam 
aijfe.  Stevsart  and  IJn'^ntT  lia\e  iimre  reeentl\'  thrnwii  cniisideralile  dniihl 
nil  these  eiiiicliisinns  I ly  liiidiiii;  that  eats  in  \\hi(di  linth  adrenal  <ilaiids  are 

entirely    reiiin\(,|    Trnm    ihe    inlliieni I'   the    nervniis  system.    Iiehaxe   like 

linriiiiil  aiiiinals  when  I'ri'jhleiied,  and  de\elnp  li,\  iiei-irlyeeinia  when  a-- 
Iiliyxiated  nr  eili.  li/eil.  |i  is  searerly  necessary  In  pniiit  nut  that,  until 
it  is  detiiiitel.\  eslalilished  li\  e\perimental  in\  esti^atinn  llnit  epiiieplirine 
may  he  diseh.  ! '.^ed  in  e\eessi\e  aiiinunts  under  certain  ennditiniis.  i|  i^ 
irral  imial  tn  a^'-niue  that  such  may  nccur  in  disease.  '/',  <  siirijntil  n  muriil 
"I    lid    'hhiiiiil   i/liiiiil    IS   iiitiin'I'i    It'll    ii'inniihil    mull  r  imif   i  n'rii  iiislniifi  s. 

The  Association  of  the  Adrenal  with  Other  Endocrine  Organs 

Wi'  ha\c  at  pieseiit  \er>  little  accurate  and  reliahle  inrnrmatinn  nii  tin 
assnciatinii  nl'  the  adieiial  w  itli  ntliei-  I'lidniTine  iii-ca!,-.  That  epiiiepliriin 
has  an    inlliiciice   mi    iiiaii>    divei-si'    nr^aMs   and    elands   is   an    lindniihted 

Tact,   lull    iIiIn  is   innre   prnhalil\    In   \»'  altriliutcd   In  an   act  i\  al  ill;r   ililluc ■ 

nil  s\  inpathet  il-  ner\c  cndiiiL;s  than  In  aii>  spccilic  relaliniiship  hetween 
file  adrenal  udaiids  and  the  ulaml  iii  (piestinii.  The  must  important  nf  iIm 
results  that  lia\e  hi'i'ii  nhtailied  are  the  t'nlhiwiiiir: 

I  With  the  Thyroid  and  Parathyroid,  ('aniinn  ,Tiid  ('.itteli.  after  em. 
firm  in 'J-  I'.radfnrd  's  discii\  iv\  I  hat  ,in  elect  rie  current  nf  act  inn  is  set  up  ii 
the  salivary  'jl.ind  \\  hen  il  is  excited  tn  ;iel  i\  tiv.  prneeeded  tn  InvestiLMt' 
the  ncciirrciicc  nf  siich  a  current  in  the  thy  mid  eland.'  I'>y  placiiii.''  nie 
llnlipnlari/alile  elcetrnde  nil  tlie  iil.ind  itself  and  the  other  on  tll<'  lieiLdi 
liorinir  sulicutaiiemis  tissues  nr  on  the  Iraidiea,  a  current  was  found  to  hi 
set  up  li\  si  imulat  mil  nf  I  In  s\  mpat  iMi  ic  !ier\  e  siipjily  ^i\  the  t  liyi'onl,  l'.\ 
iiitra\cnnUs    inji.iinn    nf    epinephi  ine,    m-    hy    stimulatinn    ni    the    crc.i! 


i  I 


TIIK   ADUKN  \l.  r.l.  VM'S 


747 


N|,liiiH'linic  iiri'vc  licl'<:rc  it  i-ciicli.'s  tlir  ndrcii.-il  -IjiimI.  This  last  rrsviJl, 
wliicli  isllic  iiiiisl  iiiiiKiitniit  ill  the  pfcscnt  .•(iiiiicction,  was,  h.iwcvor,  nut 
iil.scrvcl  wiicii  llic  IiIochI  (it"  llic  inl'ciiiir  vena  cava  ua-.  prcvcntcil  liy  llif 
ai-plicatinii  of  a  clami.  fmin  i;rttinir  to  tl'f  iMaif,  Imt  iiiiiii(<li:iti'ly  ap- 
prarc'l,  after  sliiiiulat  ion,  wiirii  tlw  riaiiip  was  i-ciiio\  ,•,!.    Tiiis  cxi.criinfiil 

t;;U.';i  alone  cjo.'s  not,  iiow  ever,  jiistifv   1| iielii-inn   tliat    tlinv   is  any 

,/,>,,/  n-lationsliii.  hctu.eii  tlie  a<lr.'nal  ulainls  ami  the  tli\ioi,l,  iM-callso 
tlw'iT  ai'c  ill  the  tli.\  idi<l  iilan.l  st  rnctiirrs  sin-h  as  tlie  iii\is.-le  lilicrs  in  Die 
l.joo.l  vessels,  wliiell  il  liyi.er>eelet  ion  of  e|iilie|.lirin..  ini-llt  alTeet.  !leforo 
any  <iii'eel   relationshii.  l.etvveen  the  \\u,  -lainls  eoiihl  he  elaiiiied  to  exist, 

it  \\oiil.l  hen ssary  to  show    that  th.'  th\roi,|  aelion  euireiit  is  ohtaine.j 

uitli  a  eoneentratioii  of  e|)ine|.hiine  in  the  liloml  h-uer  than  that  atTectinu 
the  lilooil  vessels. 
'J.  With  the  Sexual  Glands.     As  mei.iion.d  al'ove,  a  very  .lireet  lehi- 

liolishil.  exists  hetween  the  .levelopiiient  .if  the  sexual  ulainls  ami  tliat  of 
lh.'  siipraicnals.  particularly  the  .-oit-'X  of  the  -lands.  In  addithm  to  the 
L'viilem-e  ahove  funiishe.!.  it  nia.\  he  iiieiitione.l  tliat  in  hype-plasia  of  the 
.•nlreiials  eliaimes  oeeiir  in  the  testi.'les,  paitieiilarl.v    in  tleir  interstitial 


:!.  With  the  Liver,     of  the  man.v   funetions  of  this  -laml  that  whn-h  is 

most    .lireetlv   a.sor'    T.l    with    epinephrine    is    lhe    prn.lu.'tion    of    -lueose 

from  ;:ly,-„.^en     -h-  ,,!>  en.„-no|,M ,r  pr .s   .  >,.,•   , e  HiIM   .     The  in.ieetio,, 

of,.piiiephrineeausesan  immediat,.  diseha  r-e  .,f         '    an  exeess  of  l-dueose 

inin   i|„.   hi 1   that   h.vper-lyeemia    and    -l.veosuna    immediately    follow. 

This  result  is  most  strikin-  wln'U  the  in.j.cti.ni  is  made  in  -lyeojien-rieh 
,,nimals.  In  animals  from  whieh  all  tiie  ^lyen,,,.,!  of  the  liver  lias  heen 
nmioved  hv  starvation,  tim  iii.ieetion  of  laru,  amounts  of  epinephrine 
.-.-.nses  -l.veoiieii  to  ar.-umulale  in  the  liv.M  -ells  a  re>ult  wliieh  it  1- 
ditlieiilt   ti>  interpret. 

Ill  the  liudlt  of  the  fact  that  stimulation  of  the  ijre.-it  >planelinie  nerve 
,-aiises  a  demoiist  ral.le  ineieaM'  of  epinei.hrine  in  the  hloo.l,  a  natural  .'.m- 
.dusioii  is  that  the  ul.veoMiria  and  h.v  per-lyeniiia  whieh  are  known  to  re- 
sult from  stimulation  of  the  sphinehiiie  lu'rv.'  or  of  its  .-enter  in  the 
„i,,dulla,  must  he  .l.'p.nd.'iil  upon  a  hyp.Ms.-.-ivt  i.ni  of  epinephrine. 
Kvhl.'m-,.  >uppo.tin.a'  Ihiv  hyp.ith.'si.  M-,ni..l  t..  Ur  fnmish.'.l  hy  the  ol.ser- 
xati.ni  that,  affr  the  r.mi.>\ai  ..f  the  a.lr.nal  -ian.ls.  stiniulatmn  of  tin' 
splanelilii.-  ..r  ..f  th.'  s..-..alle,l  ••,|iah.-t  !.■ '■  .-enter  in  the  f.Mirth  velltrieh- 
,,„  loiiirer  pr.Mlii.-e.l  -^l.v  .-.'suria  even  in  a  ..d.v  .-o-.-n-rieh  aiuiiial.  Hut  it  is 
diflielllt  to  see  Imw  sileh  all  imp.u-tant  ph.\  si,,l.,-ie  pr.ieess  as  that  of  the 
nerve  eontrol  ..f  tin'  pro.lii.-t  ion  of  su-ar  hy  tin'  liv.-r  shouhl  he  depeudent 
oil  the  hvpei-s.-ereti.m  oi  ih.-  a.''-<iial  -iaml.  esp.-eially  sir.ee  the  epin.-ph 
line  woiihl  hav..  to  he  .-arrh-.l  hv  the  hlo.,d  arouiul  a  eoiisi.h-ral.le  pan  ol 


')| 


n\ 


1 

rnr 

1 

i*  ,1, 


!/■;■'  '^  iff''! 
V  ' '      ' 


748 


■nil      I  SIMM  KIM     ii|;i.  \N^,    lili    III  I    I'M  >-    (.l.\M 


tlic  riiciiliilioh  li(  f.iiT  it  alTiM''!  lit  till'  |il:ii-'  ii'rli  it   is  tci  iic-l.     .Mm-i'- 

n\rr,    it    li;is   ]u-iu    ^lin\\  n    lliiil    st  liiiiiial  iuii    ■  .••    |ii-r\  ioiisl  y    cut    licpati'- 

iirr\<'  ]ilt'\iis  I  jiii.iiiiil   ilic  lir|ialii-  lu'ilii-lr      ni  a   ii>ii'ni;il  animal   iiroduiTs 

liyptM'L'lv  i'iiif('ii(il\  >i-..  in  wlii.-h  case  till  IT  ciii  1"    iiii  (|iicstiiin  "I'  a  li>  pii- 

secret  iiili  111'  epilieplililic. 

\i)  dmil]!  tlie  ailieiial  elainU  lia\i'  soiiir  imi  .iitaiit  rclat  i^iisliip  tn  tin 
iierv  e  emit  I'l'l  111'  t  lie  i;l\  ein;eii,il\  I  i<-  pi  (ici -^N,  lur,  in  animals  I' nun  w  liicli  the 
aijrcnal  clamN  lia\e  lieen  reiinne.!.  ^limuialinn  ni'  the  hepatic  plexus  diicv 
iHit  pi-iMJiiiT  li\  pei-'jl\  '  1  iiiia.  I'l'i'iii  lliis  'csiili  ii  \>n\|iii  appear  that  tin 
presence  iri' a  (•■■rtain  aiimuiii  ni' epinephrini'  in  the  hlnuil  is  necessary  tin 

the  prnper  1  ralisi]iis-,i ihe  im:  \  ,■  iinpilKe  rrnm  the  s\  mpathet  ic  ner\  r 

fillers  til  the  liver  eelL  When  ihe  iieiMiiis  svslein  is  stimulated  in  such 
a  way  as  In  excite  the  e|s  ciejenulyl  ic  pnicess,  twii  erreefs  liiith  operal 
iliC  ill  llie  saiiii-  dii'ciinii  with  it'^aid  'h  ll;e  'jl  \  eiiu:euic  riiniMiiili  an 
develciped:  the  iMie.  a  h  \  peisicicl  inn  iil  epiliepll  line,  wllieli  ai'tivates 
till'  sxmpallietic  nel\e  indiii'js,  the  ntliei',  the  t  raiisniission  nl'  the  liel\e 
impills,'  tn  tie  li\er  I  .11    '  .Mai  1.  I  il  ami   I!.  I  1.   I'ean  e    ." 

4.  With  the  Pancreas.  The  i'lnictii.n  ni'  the  pam-reas  here  cnncenie.l 
is  thai  III'  its  siippiised  internal  secietinn  iVmii  the  Isles  nl'  Laiiireriiaiis. 
Since  epinephrine  readil\  prndnees  ulye.isiiria,  and  since  excisimi  n! 
tlie  pallcieas  has  ihe  same  ri'l'ect,  il  has  lieeii  naliiial  In  iinpiire  whethei 
any  relatimiship  exists  helween  the  twn  glands,  and  snnu'  iiliser\ers 
ha\e  nlilaiiii'd  n-siilts  \v  hich  they  iiileipiet  as  indicaliiiLr  that  it  dm^ 
('ertaiii  iihsei\ii-s  .•\en  state  thai  ■Jvcnsuria  dues  imt  nceiir  alter  the 
inji'cfiiin  il'  at  the  same  time  i  xliact  nl'  pancreas  is  iiijeeled.  It  is  al 
liinsi  certain,  hii\M\er.  that  thesi-  icsiilts  are  iint  t  rust  wort  liy.  'I'liils, 
remnval  nl'  the  adiiiial  ulands  iii  an  animal  siilTeriiiL'-  I'rnm  jiaiicreat  |.- 
dialietes  lines  lint  leslnie  ally  n  1'  the  Inst  pnwer  111'  lltili/inir  i;llle'ise 
dliriliii'  the  t'en  Inniis  that  the  animal  remains  ali\e.'  'i'liat  snme  n  la 
tintisliip  iiia,\  .  Iinwexer,  exist  is  indicated  liy  the  fact  that  epineplirii'i 
causes  dilatalinii   nl'  the   pupil    when   it    is  dr.ipped   inin  the   e\i'  nj"  a    p'l- 

snll    slllTerilrj     I'rniM    dialietes.    whereas    it    has    hn    slldl    I   rrecl     ill    the    nniiiia^ 
ii'iliv  idllal. 


:(t 


(  ii.M'Ti:i;  i.xwiii 
Till',  Tiivi;iiii)  AM)  i'.\i;.\iiivi;(Mi>  t;i..\M)s 

structural  Relationships 

Ti„.  tl,M-ui.I  aii.l  pnntlivn.i.l  ..laiMl-  ;itv  intiuiatclv  a>s.MMMlr.|,  iniatom- 
i,.;,llv.   ill    iiin.t    aniinaU.      Tli.   tli.rni,!    is   ,,ivs,.nt    in    all    lhr    v.mM rl.ratos, 

|.,i1  'lliP   i.arallivini.is   ,l.i    im!    ur   U■\„^,    \hv    aniphiliia.      'Vh'-    ilivroi.l 

rvists  as  INV..  lat.'lal  lul,..>  Jniiir,!  nv.T  ill-  1  larlira  In  llic  s„-callr,l  islliuius, 
Tl,..  paratiivroi.ls  a.v  vnv  .,iu,-li  MnallcT.  l.-in-  Iniir  in  nmalH-r  iincl 
,,„,„..,1  in  ,,ai,N  „M  llH.  pnstrrinr  as,„.,.t  nf  tl,r  ,l,.v,ni,l  1„1m...  TIm.  two  M|.|-- 
,,;,,'atlivnn.Is  a.v  n.nallv  iiimr  ..,•  !,•.>  nulM.l,!,.,!  in  111.,  tli.vim.l  1  ,ssur. 
ulHMcas  lln.  Inu.T  nnr.  a IV  niii.-li  inmv  l„n.,.|>  alta.-lnM  to  tlir  tli.vronl: 
in,l.v,|.  in  vnnc  aniniaU  tl^-v  a,v  ,piilr  -paralr  iVoiu  it  an.l  may  1>r 
l,„-at.'.l  at  a  .iistan.-..,  a.  in  tl,c  i„.,lia.tinnin.  .\.v>M.ry  lliyrowl  ami 
,,aratlivioi,l  lilainU  a.v  sMinrtinHs  fivs,,,!  in  tli..  liss.l.s  of  llio  .uvk.  o.' 
i„  ,li,.'a.i1.Mior  .n,..lia>tin.i..i,  ar,M..su,y  |.aratl,y,„i.ls  hoinir  .•oii.inoi.  ... 
til,,  .ahhit  a. 1.1  .'at.  a.i.l  para.  Lynn. 1  tis^n.'  li.nnu'  i-ivs.^i.t  i.i  tlu^  tl.y.i.us 
ill  :,  ,„.,.  ,.,.|it  ..r  .Inos  i  \laiiif  .  I'-rf'-iv  th.'s,.  aiiatnini.-  ivlat  ioiisli.I.s 
wc.-o  11i.il-..ll'-'lil>  \\oik...l  .inf,  tin  IV  uas  inii.'li  .•..iii'n^'  in  ill  tlu'  iiitc.'i..v- 
tati..ii  ol"  111.'  ivsnltN   rolL^uinu   niaoval   ..T  .'ii.'  "v  \hr  .itluM'  -lan.l. 

Ill  tli.'ir  liist.ilo-i.-  still. •imv  an.l  .  niln-\  ..L.ui.'  .l.Tivali.....  Hi.'  1"" 
..'laii.ls  aiv  \.Ty  .liff.nv.il.  Tli.'  paiatliv  r.n.U  aiv  .l..\  vLiiH^.l-as  an  ...it- 
uMowlli  fiviii  tlic  lliinl  an.l  r..iiiili  Liaiirliial  imhi.'Ii.s.  an.l  tli.'X  a.v  .-.m- 
,,os.^.l  of  iiiassrs  of  .•pitli.-lial  lik.'  .-.lis.  s.mi.i ihi.^s  io..iv  -n-  Lss  .livi.l.Ml 
ii|>  int..  l..l.iil.-~  ..!■  tialMvnla-  l.y  lian-K  ..i  .•onn.M-i  i\v  tissii.v  Tli.'  .'•■lis 
,-.,ntaiM  yranill.s,  s.ini.'  of  ulii.'li  aiv  ..i    a    laity  natinv.      Snni.tim.'s  .'..l- 

Ini.l-liU..   .iiat.'.-ial    is   loim.l    L.'tu.vn    ll IN.   ...•   it    ina>    l..'   .•ii.-!..s,.,l    in 

sniall  v.'si.'l.'s  .n.t  iinlik.'  ll."-.'  "f  'li.'  tli.M-.ii.l.  aitliou-li  iisiiall>   .•..iisi.l.v- 
al.ly    small...'.      Tli.'    I.I0...I    n.-.U    aiv    .■Mivi,i..ly    iii.iii.T..ns.    an.l     foim 
sinilslik..   .■ai.illari.s.    ^^l^l.■ll    .-.■m.'    into   ..l-s.'    ivlat i..n.-Iiii.   N\itl.    tlu'    cpi- 
tlu'lial  (••■lis  of  til.'  iilan.js.      N.i-v.s  aN,,  ar.'  al.ini.iant   an.l    pass  l...tli   to 
llu-  vessels  ami  t..  tin-  s...Mvtiii-  .•.lis,     Tli.'  h\>nn\  vss.'ls  a.v  .I.tIvmI  from 
the  iiit'.'vi.n-  lliyr.ii.l  ait.'r\-. 

The  tllMvi.l  is  il.\.l..p.'.l   !..v   iiiim.'.liate  ..lituivwtli    f.-olll  the  ent..<leiiii 
liniiiu'  the  ,1......  of  the  pliaiviix,  at   a   l.\.'l    l..'t\\ee..  the  li.s'    a..,l  se.'.'H'l 

Jn-a.iehial    i>.....-hes.       |{..pivs,.,it.M|    at     liisl    l.y    a    soii.l    .■..limin    of    .■•■iis. 

ther..  ve.'y  s ..-.•ni-s  a  .livisi.m  at   ih.'  l.iu.r  eml   inln  tw.i  lat..fal   p.).'- 

749 


i  Ic    I 


i 


i '  i 


•I   ,  I  I 


T.'iO 


I  HI     I  MHii  i;iM     ii|;i.  \s-.   Hi;   hi  I  11  I  >■-   i;i  \Mi^ 


tiiill^,     ;illll      tllr     iM'rjilul      M.li.l      ■■nluillll      I II     s      llnl|n\\c,l      Mill  'I'llC      I  W  '  > 

l;il('i':il   1ii;iiii-Im'v  oI    lln-  orii;iii;il   roliitiiii  dixiilc'  ;il:;hii   .iiicI   :iL;,iin   mi  us  t" 

I'liMli    -.[   s\-.1riii    ..I    Imlluw    tuli(>    !1 1    \\\\\\    i-pitli.'liiliil.      Tin  sr    ;i  I'l  ri'w  :i  n  I 

Iii'i'iilllr  cut  ll|)  Nil  ;is  111  I'lillil  tllc  clii^r.l  Msiclis  rlul  |-,-|ct  I'I'InI  ii-  i,\'  \\,,- 
'jhiliil.  |-!,h-ii  \  I'-ii-lr  i-«  liinir  nr  Ifss  s]  liir  ii  lii  hi  i  III  sli;i|ic.  :iiii|  li.'ls  'm 
li;iM'IIHIll  111!  liiliLllir,  lill)  itv  \'.  .-llis  ;lli'  t'i.ri||ii|  li\  ;i  I.IViT  III'  c  pillirliii 
i-(ll>.  \\liii-li  liiii\  111'  cnliliiilKil',  riiliir,-|l.  or  ll;il  Iciii'il  ill  Nliiipi'.  \',;i'-]i  \  csirli- 
is  lilli'il  witli  liic  vii.imIIiiI  mil. lill  material.  \\liii;li  i-~  |pci-uli,-ir  in  run 
tainiliL;'  iniliiic.  ainl  lirtwiTii  llic  \r>ii-lis  In  a  laviT  ■■!'  i-'Uiiici't  i\c  tis^iii 
nfli'ii  i-niitainiiiL;  small  rclU.  ^imr  nl'  wiiirii  arc  imt  iiiiliUi'  tlmsc  ciT  lli. 
paiatli^  iiiiil.  'I'Im-  ciiiiiiccti\c  lisNiic  also  cnntaiiiN  lli,'  liluinl  \csscl-, 
whicli  arc   \civ    iiiiiiiciiiii>      iiMJci-il,  tllc   llivrnicl,   in   iiiupiu  t  imi   tu   its  ni/c. 

iccci\CN   mwic    tiiaii    li\c    tinier   as    nnicli    lil I    as   the    kii|iic\N.    tl hi' 

tissue  thai  Niii  pa^Nc-.  it  in  tlii^  icjaiij  licini;  the  mciiiilla  nl'  tlic  adii'iiai 
jla"il  M  •■  \r.r2r  'Jll  ,  Till'  iiciAcN  ari-c  rfnlii  Icitli  the  \a'^u>  aihl  tin 
N\  mpathet  ie  system-,  ainl  have  lucn  iiaceij  1,1  ilic  -.cei  et  iir^'  cpiilielial 
eelU      The  alii'xc  ii',ciipt  inn   app^lc^  Id  a   >liielly   iiniiiial   ulainl. 

THE  THYROID  GLAND 

Condition  of  the  Gland 

In    tile   ciii'mIc'1   cninmiinitics   i.T   the   (lieat    Lakes    Hasiii    <>]    this   cimti 
iieiit.  it  lias  hecn   fi.iiiiil  thai  in  innsl   animals  the  l!i\  i"!,!   .jlaiid  is  imn'c  ••' 
less  ahiinrmal.      In  ('Icxclaii'l.   \'<v  c\aniple.    Mafinc   has   iminil    this   In   he 
the  i-asc  in   \\<11   n\ei   !li>  pef  cent   <<\   the  ihejs  lir(M|eht   1u  the  lahnratnry." 
The  (Mirilii'iii   Usually   'jncs  uiiijci-  the   name  oi'  simple   eciitcr,   whjcli   in 
i-ln.lcs    all    thyr.'i.l    eiilareements    c\cep|    tlms,.    ,,]    c\(i].hltiaimi<'    u'oiliT 
In   man   tlic   cnitii-  niieinatcs  usually  almul    the   a'jc  nl'  ailntcseiiice  ;niil 
mine  I'll  ipiciitly  in  uiils  than  in  Ihivs,      It   ma\-  snmctiiiies  pass  i>\r\-  intn 
the   cNnplithalinie    type,      'i'lic   exact    iiaihnhicic    cliainjes    in    tlic    i;ipit  ciiii- 
clanil   \ar\    with   tlic  species  nl'  animal   ami   with   the  ijiiratii'ii   nl'  the  ilis^ 
casi'.      In  man.  liesiiji's  the  e\stie  m-  cnlhiiil   e'>itei-  .-ni  ailcnnm.-itiiiis  Ivjic 
is  \{v\    enmmnii  altiiniieh    rare  in   nther  .■niimals. 

l-'fi.m  thi'  nuniei-iiiis  (ihsiT\  ,il  imis  th,-ii  iia\c  heen  m.nle  nii  the  chmils  nl 
(liiiiicstic  .iniin.als.  it  li.-is  liecn  elcaily  cstaMislicil  that  ilie  \ciy  caflicsi 
siun  nl'  Igniter  is  a  iliminiitinn  in  tlie  iniline  ciuitciit  nt'  the  ijland:  In! 
Iiiw  eil  liy  ;in  iiicrc^ase  in  the  cpithi  'i.il  cells  a  ml  in  the  hhiml  supply  !iml  a 
'lecrease  in  tlic  cnllni.L  Siieh  Ini/n  iiilii\ni  \n;\\  \ic  iiiiluce.l  in  what  ic- 
mailis  at'tei-  |-cmn\;il  nt  a  laiL'e  pail  nl'  a  mniiial  chnul  '•niiipciisatnix 
li\  pel  piasiii  .  ..i  i;'  ,1  siiiiilar  ..j.i  rat  i.m  i.c  |ii  ri'm-mci!  early  in  pnunalir;  . 
tjie  yniiip,'  win  11  hnni  \\ill  he  I'liiiml  tu  haxc  hy  pcipl.-istie  th,\  rniijs.  A 
iTrltiin  ilc'.'rce  nl'  !'\  pi-ipl;isia  exists  as  .-in  accniiipaniment   n|'  pfcirnaiu'y. 


■1111     I  H\  l;iiiii    \Sh   r\ii  \ni\i: I  \M'^ 


T.'.l 


.,,,,1  i,   ,.„n  li.'  i.i-.m1ii.t,I  ii,   i,uriM:!l  iiniiu.iU     f;!  i  t  i.Milail.N    rMl>'    l.v  l.liiclir.' 

,1„.,„  „„  an  .s.MsMv,.  i,ir,-,t  .li.l       .urlaiil  ,,I,-«.t\  atioiis  licariiL.'  •.„  tins 

l„,i,„    |,,.,vc   Imti,    liiii.l.'    I.v     Miiiiii I'i-""1<    "-"'I''    '"    ^^''i''''    ''    ''''^   '"'".'" 

fniliul  llial    111.'  s.,-ralli'.l   .•an'innina    llial    .lev. 'lops   "  l'"li  lli«'   li>li   l^-'l''    '" 

|,.,t..l„Ti.'s   aic    iVd    villi    niiMiilalilr    I I    an. I    ..v  nviuu  ,l,.,i,    is    nally    a 

ivpiral  In  iH'i|.lasia      In  its  M-.n.i  -ta.^.'  this  ,1,-n -li-i'^  into  uliat  is  known 


■I       I 


•^s. 


:•«  %..<'' 


\> 


:i/i 


^''' 


.-'4-.^-.. 


V„ 


\       ...    ^-^ 


!: 


as  ci/ZoiW  ;l'>llir   \\\\\vh    is    |n(iilii 1    I'.v    a    .1, 'position    m'   ,-nll..i,l    mat-'rial 

l...tw..cn  tlH'  row  of  rcll--  Ml  as  to  .-aiis,-  an  ..|„Miiiiu  out  a-ain  n\  llir 
v.'sicK's  I  !•'!•;■.  lit:'.  .  \utli  a  .•nnscMUcnt  lrn.l.-ni-\  tn  a  vcvtM-sinn  to  tlir 
n„n..al  histolooi.-  si  ni.'t  mv,  so  Tar  as  tliis  is  |.ossilili-.  Tin'  vrsi.-U's  in 
ii 


>u,-i,  a  ^land  alv  of  .nonno,;,  ,i/,-,  an.!   th,-  iniin-  .  pitb-tinni     !oxv  ..uLi-il. 
or  alnnist  Hal  in  stia|n'. 

Till'  oulslainrni..'  .-liarai'tcrisl  i,-   f.'atiiir  of  tlic   rolloi,!   niat.nial    is   tlial 


!l.f 


r:  ^T 


•nil     I   \|Hir|;|\|     (ii;i;\\-,    ni;    HI  i   II  I  »   (J  Wn 


■:l, 


it  I'liiiliiiiis  iiiiliiii',  wliicli  exists  in  niniliiiMl  inn  willi  ;i  iiiiii|ii'iil('iii   iiitm'; 
ciKPils   liMsc,   iiinl    is   iiMiiilly    fjiliccl    imlollix  riii.      In    the    >:\i\\\i\    ilsell    tli. 
ici«li)tli\  rill    iiiii\     l>i'    in    iMHiiliiniitinii    witli    prutciii,    rotiiiins;    iddidli.viH 
V'liiliiirm       !■;.    (',    Krniliili"'    liiis    rccciitiv    sliccf'cclcd    i       isniiilinK    n    ])lii. 
n-ysliilliiic   siilistiiiiii-  111'   pel  iVi-tly    c-iiiist;iiit    coinliiisili  mi    Mini   cciiitiiiniiiL: 
nvrr  tl((  jKi-  (Milt  lit'  iiiirnic.     It   is  r.illnl  alpha  iinliiu ,  ami  lias  lii'cii  iili'iiti 
lii'il  as  an  imloli'  riiiiipoiinil.     In  i'\tri'iin'ly  iiiiiiiitc  ilusaj.'!'  it  irri-atlv  alVcri^ 
tlir  ('ii(ii.'.\   iiictaliiilisiii,  ami  is  saiil  to  imliH'o  syini>tiiiiis  liUr  rxiiplitlialinii' 
U'liitcf.     Its  tliitapi'Utic  value  in  eases  iif  tli\  loiil  delieienev  is  iciiiarkalile 
Kemlall  lielicM's  tiiis  siilistMliee  tii  lie  tile  iietive  emistit  Uelit  iil'  the  tli\  liiiil 
ami  to  lie  assueiateil  w  illi  tlie  met  a  In  il  ism  ol'  amino  acids.     I'm-  one  lliiii'.:. 
when  il   is  liiveii  almie  no  ehaiMje  ueeuis  in  jiiilse  rate,  whereas  if  aiiiiii" 
aeids  are  yixen  alonu'  \xilli   it,  tiieie   is  aeeeleralioii. 

The  import  a  nee  of  the  relationship  lietweeii  the  t'unetion  ol'  the  lh>  mi'l 
and  the  iodine-eonlainiiiL'  material  is  indicated  liv  the  chaiiL'es  which 
occur  in  the  i>ereentaLre  of  iodine  in  the  inlands  under  \ar.viiii,'  condi 
t  ions  of  activit  \ .     Marine  oiiserx  ed  that  the  amount  of  iodine  is  iii\ crset) 

lnoportional    to   the  dciir f   hyperplasia   of  the   '-dand,   and   when    tie 

hvperplastie  condition  heeomes  fully  de\clopeil,  si-areely  a  trace  ol 
iodine  is  eoiitained  in  the  Lrlaiid.  Later,  \\hcii  the  hyperplasia  yiM- 
|>lace  to  colloid  L'oiter,  the  iodine  increases  auMiii,  Imtli  ahsoliitely  and 
relatively.  Miireo\er,  it  h;is  liecii  found  tli;it  if  iodide  lie  adininistered 
to  an  animal  sulTeriiiir  from  li>  perplasia,  the  hyperplastic  condition  vcr\ 
ipiicklv  disappears  i  Kits.  Ilfji  and  the  animal  heeomes  iioriiial.  Thus,  in 
lirook  triuit,  the  poor  nutritive  conditioii  of  the  (isli  when  hyperi)Iasia  ha^ 
developed  can  he  immediately  reiiiedied  l>y  plaeiiifr  thein  in  larizei'  (plan 
lilies  of  ruiiniii!.'  water  or  hy  addinjf  small  ti'aces  of  iodide  to  the  water 
The  administration  of  small  amounts  of  iodine  as  in  ordinary  salt  frotii 
salt  deposits  also  incvcnts  <:(ii1er  in  I'arm  stoids.  lliis  haviii<i  liccii  first 
liote(l  in  the  State  of  MieliiL'aii,  where  inior  to  the  discovery  of  sail 
deposits  sheep  lireediiii:  v\as  an  entire  I'ailiire.  The  importanee  of  admin 
isteriiifr  small  doses  of  iodides  to  sidiool  children  livinir  in  i;oitrous  dis- 
tricts has  recently  lieeii  cmphasi/ed  hy  Marine  and  Kimhall.'^  As  small 
a  (lose  as  0.(101  ij-m  .-it  weekly  iiilerxals  prevents  iroiti'i-  in  puppies  sus 
ceplilile  to  it. 

Experimental  Thyroidectomy 

A  correct  interpi  etation  of  the  functional  chanires  and  svniploms  which 
follow  upon  partial  or  c(>mi)lete  removjil  of  the  thyroid  iriand,  or  from 
its  disease,  has  proved  a  very  difiicult  pmlilem,  parll.v  liei-ause  Mifiicieiil 
care  lias  not  heeii  taken  to  note  how  much  iiaralliv mid  tissue  was  re- 
laoved  aloiiij  with  the  thvroid,  and  partlv   hecause  the  I'aet  has  lieeii  over- 


IIM      TIIMJUII.     Wli    |-\l!\TIIVl;nlh    111    \M>S  <■'•• 

1„„K..,1    lliMt    tl„.    Hr.Hs    ,.ro,|u 1    In     llnP.i.l.-tn.MV    .n,l    ,.Mrall,>  n.i.l- 

,.,., V    an.   ..l-t,.M    N.Tv    ,l,ir.-Mt    ,n    animaU   ul'   ll,.    si Kim,1    at    .1.1 

iVn.nt  a->     SpraUii,..  ....,M..all>  .  il   .Ma>   Ur  sai,|  llial   ll.,.  intlu,. 1   Hi" 

,„„,„l,v  ,.„i.l  i.  rn.M.-,|  .nainl.v   on  tlir  ihtv.  r.-.,lr,.  aii.l  -hiIv  t.,  a  s .n.l 

,,v  ,|,.',M-.T  ..h   llir   .iirlal.nli,-   rnncli„ns,   u  lirrras  llir   ivn.tm.   is   tl...   .-as.. 

ui'll,  tlir  llnn.i.l.  its  main  HT..-t  Lrini:  .m •tahoiisin,  althon-h  il   l-i-l' 

;,1,K    also  (.N.T..is,.s   a    s ,n.lan    HT.H    on    tin.    nrrv •nl-ns,      Moiv   sn 

,!,,,■„  in  tl,r  .as.,  of  anv   oHht  nnlo.-nn,.  nr.^an.  our  Knoulr,!.,-,.  i.-nnur.' 

,,,„   f.UiHion  of  tin.  tl,>toi,l   l,as  W.u   U-Vno.,!   Uv   ..hnn.al   .■siMTirn,-,  ami 

,1   is  ,iil-|irn!'    tM  >a>    ulu'thrr  111.,  .-liniral   nv   111..  .AiM.niih.ntal    IImmI    !ias 

,.,,n1rilMit..il  tli(.  -T..al.r  anioinil   of  inl'oiinat  ion. 

Tl„.   n.snlts   „f  ..NiHrinnntal    ..xtirrati f   Hn'   lli.M"i'l    ^  ■">    ^n-ronl- 

ii,.,.  to  111,,  au..  of  tin.  animal,  au.i  fmin-.n!  ly  Ihrv  nvr  l.y  iio  niraiis 
,„a,.k,.,l  i.rovi.l..ci  snfli..i..n1  paiatlivroi,!  tissue  lias  Ur..,,  uiMlania.j..,l 
Tl„.  svinploMis  ar..  in  urinral  tirn-kmin-  ami  ilninir  of  Hi.,  skiii,  uitli  .i 
,,,,.|..n..v  t.i  a.liposilv  ami  a   loss  of  ton.,  of  tlie  nuisrl...     Tim  lio.ly   t...n- 

,„,,,,tuiv   is  low   an.l   tin-   s..Nnal    fmiHions   1 .iir   sulniormal.      NcTVoiis 

svn,|.ton,s  in  tin.  ,rm..-ti..n  of  ni..ntal  .Inllm.ss  ami  l..tliar-y  an.  also 
usuallv  |.ns,.nt.  Snn/n-al  n.|llo^al  of  tl,..  tli>n,i,l  in  man  iH..Ml,m..s  tli.' 
,,„„|it'ion  knoun  as  n,.  In  .,;„  .Ir,nnli,nr„.  Tli.-  symi-loms  may  lirsl  ..I 
,„  ,„„.„„„.  „,,,,,.,nnt  a  f.'w  .lays  aft..r  tli..  ..pc-ntion.  o..  tli..y  may  ninain 
|;„,.nt  for  vais,  ami  tli..ii  .l..v..lop  so  as  to  pn^.h.....  tin'  .•on.l.li.m  km.un 
■,s  mvv...l."mM  Winn  n..iv,n.s  sympt.ims  an-  pn.minnit  ni  ..a.-ln.xia 
s,,um"ipiiva.  it  is  nsnally  tak..n  as  ..vi.l...i....  tlial  an  ..x.^.'ssiv,.  aiinnnit 
„,■  ,,.„atl,vnml  tissn,.  lias  l„.,n  ,l..st  my,..!.  Ko..|i,.r  stat..s  that  att..i-  ..oin 
.IH..  k.ss'of  11...  tliynii.l.  lit-.,  is  impossil,!..  U-v  nu,vr  llian  s..v.ii   v.-ais, 

(■tTi...ts,  at    l.'ast   on. .-fourth  of  th..  or«^an 


I' 

ami  that   to  iin.\(.nt  ultiiiiat..  il 

sh.inhl  h..  hit    inta.-t. 


t  f 
[ 

I  ■ 


Disease  of  the  Thyroid 

Th..  sM.ipt.HMs  ..f  .lis,.as...l  ....n.lili.Mis  ,.f  Ih..  thyi-oi.l  may  1...  int..|- 
pn.1o,l  as  tin.  <...iis,..,H..n..."  of  iii.M-..iis,.,l  <..•  .liminish..,!  fnn.-ti..nin-  „{  llic 
„|„n.l.  S,.m..tim..s.  l,o«,.v..i.,  th..  l..ss  a.'tiv..  .Jaml  is  n-ally  im.n.as,.,!  in 
hulk  this  in..|...as..  h..iim  .-aiis..!  hy  th..  a.-..niiinlati.ni  in  it  of  N.'iy  Imi-u.. 
,,u.„ititi..s  of  ...ilh.i.l  mat..iial  a.....niipani...l  hy  an  attonuat...!  ..on.litn.n 
„f  th..  v..su.ular  .-..lis  ( s....  pa-.'  T.'-l..  Wh.'ii  th..  ulaml  is  atn.phm-l  at 
|,i,-th  th..  .•on.liti..n  of  inlniiMii  s,,un  h....oin..s  ,h.M.l..p...l  i  Kil-'.  I1>-»  .  'I  h.' 
,.h,u..-u-toi.istic  tVatm.os  of  cn.tinism  -.m-.   iV^   An  arrest  of  -an.wth.  ..sp,-- 

,.ially  ..!■  tin-  ski'lHon,  a.-.-.mipiini.  .1  hy  ii mph't.-  ossith-ath-li  .if  the  h.in; 

i„',n(".s  ami  railuit-  of  tho  fo„tan<.!h.s  of  th..  skuH   t,,  clos..  pn.p,.rly      tli' 


I'lHir  .l..vi'l.ipm..nt  of  th..  nnis..|ilar  syst.-m. 
,-,,mlition  »<  tho  skin,  so  that  it  is  yrllowis 


An  nnhcaltliy,  iliy.  swollen 
h  in  e.ilor.  tin'  fa.'..  li(.insr  pill.. 


u   h 


if  t? 


\  : 


&  j    i 

fc    ■     ' 

1  i  ■;' 

'"' 

f  ■ 

1 

:  '■      1. 

':  ■    i 

i:.4 


I  III     I  M'(M  i:'M    ui!i,  vs  -.   (.i:    I'i  c  1 1  I  --   C.I  \\ii~ 


iilnl    pUlVv.       li'     An    ;ilihiifiii:il    i|i\rlii|iiiiriil     nl'    the    cdiii li\c    t'lNsiu- 

ciilisiiiir   ji    sliaiii'lo^ 
s\\  ollcli,    I  lir    liah'I^    ;il 
Till'  liciv  llll^  -.\  sli  111   ; 
IHllirit  \    (ir  ii\  cr,  I  III' 
iiliiiti^in   Ih'Iiiu  <''iiiiiii 


■Mhililiipii    nl'    till'    >urr.ii'i';    llir    iiImIiiiiii'ii    i>    ;iI«,'i\-, 
i|    r.rl    iiii'   ■>li;ilii  li^>,   ,'ilii|    till'    hii-i'    c|i'|ili'>M'i|.       I 'i 
Ui,  t'.iils  to  i|i-mIii|i  |ir..|ii'rl\ .  mi  llmt   ,'it   tlii'  iiL'i'  nl 

llihl    irlll.'lih^    llki     .'III    ilil.'ilit    ill    lli^    llli'lit.'l!    Iil'llll\  liM 
111.        Imlrril,    till'     H  lliili'    I'llllil'al     |lil'tl|li'     i^    s(i    I'lllil 

ai'Iri  I--I  ii-    lliMt    iiiii'c    lia\iii'^-    ^ii'ii    a    I'a^i'    i lir    i-aii    I'ail    al't'Tw  anl    In 


I     it  I'.l  I     M    1|„, 


,  I,;. I.    n...  11  .iinii'i  »itii  tliM'.i.l  iMi.iu  -I. 111.  I  ii.,  1.1U  I" 

t.'.t       ( i'.i'i.  lit  i.l    I  'I.    S.  J.   \\'«  '.-n  r  1 


ll'cii'jlli/.r    till'    llislilM'.       lii'Nl.lr-.    lirill'j'    ilui'    '(I    full'.'i'llital    allM'Hl'C    iit'    til' 

tli\i'iiii|  I  spcii'ailir  Ivpi''.  ri'i'tiiiiMii  may  also  iifi'iii'  as  a  ri'siiit  iil'  tjiiitriiii^ 
tl('H''ii''i"itiiiii  I'l'  llii'  '.'lain!.  Tills  I'lii'iiw  till'  siii-alli'tl  I'liilt'iiiic  v.'ificty  nl 
the  (lisi'asc,  ami  is  nmn'  (•(iiiiiiiuiily  si'cii  in  ^nitiniis  ilistiii'ts,  lieiiif;  lint 
iiit'ri-ijiiiutjy   a-siiciat''i|    ■.'•  iUi   ijis.-a-''   "!    llic   j>af:i!!iy!'i>ii|,   in   v,iiii-li   f.-iM- 

till'    llt'lMlllS    sylll|ltlilllS    .'III'    M'l-y    |ll'n|||i||l'llt. 

Atiiipliv    III'  till'  lli\iiii(l  in  adults  i-aiiscs  tlu'  clinical  conilitioii  known 


Ift 


Mil      IMM.'olh     Wli    rM;\lllM: i.l   \M'>  '■'•' 

,is  „H/M./.  „/.;,  iili.l   li.'i'-  a'jaiH   lli'-  ^^lnlltoMl^  .H.'    \>i>    .-liiu  ii.'I.Ti>l  ii-   '  KlU 

1:1;,  'I'll.'    >Klll     ■>    ih>     .illil    ihirL      will'    n    i|.'|">lli ,M,llUri-li\r    Ilssll.- 

nrifll    .•nl.lMill'.llU'     I'.it     il.     i'-^    .|.-,'|i(l      l,l\rl,  ;     til.'     lillll.U    Mil. I     IV. ■!     Iir 

illisli;i|.rl>-.    III.'    Ilj.-    llil''K    nli'l     \\<v    I. .HUM.'     ,.  .111.  w  liiit     .hlii  1  ■-"■■I .    -1    lluit 

■  vli,.,,  tlir  p.  i>..h  iiiLiiii-iv  :..  -i"'i!--  <i  •ij'l"'"^  ''^  il  ''"■  i"ii'-;'""'  \*'i''  '"" 

|.||.,^,.    f,,|-    III,.    MM.Mlll  -.    111.'    Ii.l.l     Inlls   ..Ml   ;    ih'l''    i>   :i    l"V.     I..»l\     llMl|MT.IIU|i'. 

all, I  il  ran  l,i'  sii,,\\  11  lli.it   lli.'  -  lu  i'j\    iiLtiilinliMii  i-  •jnatly  .|,'|,n-.M,|,  .•in.j 

lllilt    il    llrfK'ich.'V     1,1'    .i\\Ulll    l>    l,.lll'J    .•..llMllllf.j.        M     iv    saill    III.      IMTMIII    <-A 

Im1>i'  11  1;iil:«'I'  i|u;,iitity  "I' Miiiai    lli:iii  an  ,tr.|iiiiir\    in.li\  i'luiil  witlMMil   tli. 
.|i'\rl.,|ilM.lil    i>\    vI.VcmMII  i.i.   I, 111    iIm'  .l.|ii.>>ii.ii    ,.|'  llii'    liU'talMilic    I'lllM-ti.in 

.■alls.'s  III,'    pall.  Ill    I..    Ilk-    S"iiiii'jl>    "I     1 1.    Ill    «|.il,     ..r    ulii,-li.    li.,u.'\,T^ 

III..   I,.i.|>    \\.i'jlil    iiia.\    ^l.'ii.|il>     in.T.aMv      'I'll.'    -iMial    inn.-t  i-m    Im in    > 


il 

i 


1-1. .v.,    Tin,  r-l..|t 


,l,'|iivNSfi1,  .-mil  111. 'IT  is  iii\..lv.iiii'iit   iiT  111.'  ii.TV.Mis  ^yst.'iii  a^  sli,.uii   1.;. 
iiiciital  ijullin'ss  .•111(1  Icttiai-L'.w 

Alllioiiu'li  till'  lliynii.l  ■.'iaii-l  is  mih-li  al  1  ..|iliii-il  in  iii>  M''l<lii,i,  s_\  in|it.iiiis 
tli.nt  !ir.'  M'l'v  similar  may  .-lisn  ...'I'lii-  wii.'ii  lii.'  ulaii.l  is  ii!,irm,>iisly  .'ii- 
l.'ir;.'.'.!.  As  ali-f.-iily  .'Xplaiii.'il.  lii."t'M'i\  llii^  iiilai'jfimiit  is  ,liif  111. 'lil^ 
t.i  ;iii  a.-i'umulali.in  o!'  I'.ili.ii.lal  mati'iial  an.l  is  i-,'all\  an  alr.iplii.'  I'on- 
.|ilii)ii.  A  p.'ili.'iit  siit'f.'riini'  rnnii  .iiiliiiii.'  'j.iiliT  may  al  liisl  I'xliilnt 
vymptiims  wlii.'li  aiv  ilsiiall>  all  liluilr.i  t.,  a  li\  p.'r>.','i'.'tinii  ,,f  lliyn/pl 
iii:i1cri;il    iiit.i  tlic  lilood   (llic  syniptimis  will   I..'  .|,s.'rii>.'.l   iiiuiu'iliatfl\    . 

,         I       1      ,       ,.     ,  1  .r^'-         111",',-      *il     -  .    ...>|t.|!"-      ""'       ,il>lil,-.i      lllii^l'      .,('      Ill  V\.'(|. 'Ilia 

Il  is  con.'lml.'il  tli.'it  til.'  iiliovf  .'nn.litimis  ar.'  iluc  to  ilflicioiicv  ..1 
tlivroi.l  fiiiicliuii,  111-  hiii„itli!irni,lisiii.  Iitvaiisc:   ,1  j  tli.'  •rl.-ind  is  jitniplii.'ii. 


ill  ill 


iv  i'i 


a  I 


■i  * 


i 


1.1] 


i^, 


Vtll 


III     I  MMii  IIIM     iiUiivs-,    nil    111  (II  I*-    III  \\|i< 


mill  I '_' '  >ltliiltll'  ^\  liipliilllN  III  llliiv  rxliiliilnl  li\  llii'  rliliii-iil  I'liliilil  iiiii^ 
••nil  lie  |ii'im|iii'('iI  i'\|M'i'itii)'tiliilly  liv  llic  rciuinal  nl'  llii"  uIiiikI  ill  iiiiiiiiiils 
lly  iilisiM'N  iitii.lis  nil   fhi-   rffii't    of   ii(liiiiiiisliiilii>ii    nj'   llixioiil   r\trm't    I. 

I'M'tilKHIS  111  IIIVM'iIi'IIImIiHIn  pIltii'llN,  |lln||||l|  il  llll'l  ilil  ,1 1  inn  nf  till'  *>lll|i 
liilMH  ll(•l•ll^^.  \\liii|i  i-ritiiinlv  sinjL'i  vis  lliill  tlir  lr:il  r.Milsc  i>  llir  ;ilisi'ii''< 
•  •t'  nil   inlrrii.'il   sirirliiiii.     'I'lirri'    is   |irnli;ilil\    iiiitliiii','   iiMii'i'    sti'iiunir    In 

till'  utinli'  ilnliiiiill  nt'  I  lirnipt'llt  irs  lliail  lllis  rllVi-t  rimil  llic  iiillllillist  lat  iciM 
nj'  lliviiii.l  lAtriii'l  or.  iiMiii-  sii  still,  nl'  al|>|ia  imljiii'  IT  ilir  tiratiiH'iit  is 
staltnl  rallv  tiliill'.'li.  tin'  rnlilHills  I'liilil  riolii  liiilli;  an  ill  i|r\  <'ln|it'i| 
iiiint  i|iiirl\l\  ratrlii'>  n|i  with  cliililirn  nl'  Ills  iiw  ii  aui'  ami  l>rciiiiii'>,  jn 
t'\ci'y  res-icft  iiniinal.  I'lviii  il'  lliis  tiiatiiiriil  is  iml  iin.l.i  laNiii  until 
till'  r-Iiilil  is  M'M'tal  Mill's  III'  au'i",  it  is  iniiaikalili'  Imw  i|nii'kl\  llii'  lii-iirli! 
Iiia\  sliiiw  itsfjt'.  In  ii'\  M'lltiiia  aiiil  rai'lii'xia  slriiiiii|iii\  a  alsn.  tin- 
s\  inplniiis  M'ly  i|uii'kly  ilisa|i|ii'ar  ami  tin'  prisnii  liii'oiins  |iiirccil\    mn 

null   by    till'   ti'ratiiH'iit.      In   all    llns iiiilitiniis,    Imwrxn',    llii-   tliymiij 

cxtrnct  must  Ik-  ailiiiiiiistnt'il  cniitinunusly  in  nnln'  tn  iircvrnt  tin*  I'l'aii 
pi'nrniicp  nt'  syinptnins. 

(Jlliti-  (listinrt    riniii   till'  alwiM'  dcMTilit'il  cnnilitidlis  nj"  liy )>ntliyrniclisMi 
ai'c  tliDSf  pinilr     i|  hy  all  I'M'i'ss  iif  tliyi'niil  aiitiH'iiiil  ill  tlic  lilmiil,  naiml> , 
hiipfrllnirolillsiii.     Siidi   a  cniwlitimi  i-aii   In*   prnilm-cil  ixpi'iiiiii'iitiilly   in 
iiiii'inal  niiiinals  liy  tlic  ailiiiinistratinn  nf  tliyioiil  cxtfact  nr  alphn-iiidim 
(Kcndain.     Ill  man  lai'i.'t'  iloscs  arc  snnn  fnllnwrd  by  u'rcat   (pru'kf'iiiii'j 
nf  tlic  pulse  with  snmc  irrcmilnrity,  flushing  nf  tlio  skin,  iiii'i'oasrd  jn'i' 
spirnfinii.  trciimi'  in  tlic  liiiilis.  cmai-iatinii,  and  maiki'd  ncrvniis  cxcila 
bility.     Alini",;  with  tliosr  symplnnis,  mi'talmlii-  iiivcstiiratimis  liavc  slmHi' 
that  the  ciicfi.'y  mitpiit  pi'i-  si|iiai'('  ini'tiT  of  sui'fiioo  is  (iicatly  inorrasici 
bi'iiisr  si'iiH'tiiiics   nral'ly   ilniiblcd;   that   tlio   nitrodoii   rxi'ii'tinn   is   om'is 
sivo;   and   that   alimi'iitary   i;lyrMisiii'ia    is    xciy    rnmiiinnl\    pnsrnt.      'I'lir 
body  ti'mi)i'raturr  is  nnt,  hnwcNcf,  as  a  iiilc  incri'ascil,  lici-aiisf  althiniuli 
mctabiilisiii   is  f'Xi'itiil,  \  d   heat   Inss  is  i-nri'cspnndiuttly   iin-nasod.      K\ 
nphthaliiins  is  said  tn  di-xrlnp  viM'y  nn'asinnally  altir  sncli   adtniiiisti'.i 
tinii,  but   this  is  liniibtl'iil.     Lastly,  tlirrc   art'  usiial'\    cliu'i'sl  ivi-  distinli 
anccs,  althniiirh  thi-  apprtiti>  is  likrly  tn  bi'  im-ri'asi'd.     Tlu'  pulse  is  c|nii'k 
cued  after  adiniiiistratinii  nt'  alpha-indinc  mily  whin  prnti-iii   fund  is  aUn 
taken.     This  is  lnlicvcd  by  Kendall  tn  be  due  tn  the  assnciatinn  betwerii 
the  thyroid  linrmniii'  and  the  metalmlisin  nl'  the  aminn  acids. 

The  .symi>toms  fnllnwinu'  the  injcclion  nf  the  cxtiact  are  very  siniilai 
to  those  of  the  disease  known  as  i  idphlliaJinir  i/oiti  r.  Fndced,  the  symp- 
tom.s  arc  so  much  alike  in  the  two  cnnditinns  that  it  is  soarccly  neccs- 
sarj-  to  describe  tlinn  siicciaily  i'nr  the  disease  CAi'ipt  In  moniiuri  tha: 
the  exophthalmos  is  much  nmre  likely  to  be  present. 

IJke  simj)le  sfoiter  this  vaiiety  is  frnm  tliree  to  fniii'  times  more  fic- 


Till      rilVHiilK    VM'   lAIIATIIVKIiin   lil.ANPW 


7:.7 


,|U.-iit  ill  vM.iiuii  lliiiii  III  iiifii,  a  liKl  of  HiKiiirtfiiiHT  wln-ii  wp  m-atl  llu' 
r\iiliii<c  nf  iisNiifiatinii  lift wt'i'ii  th(>  tliyiiiiil  tfliiml  iiinl  tin-  iffiu-nilivc 
(iit'iiiis.  Il  is  Miid  fliat  tlio  (lisrHso  is  usually  foiiplttl  wiTli  |MTsis|i'nr«'  of 
III!'  tliviims  ifliiiiil.    Till'  »li.Moi.l  ula'i'l  i"  <'\<'plitlialinif  litutcr  is  (■iilaru«'<l, 

> •tiiiifs  ill  Idle  lolic;  il  is  lianl  aiitl  l»il|iy,  nnil  <iii  H'lsi-ulfalidii  a  niur- 

tiiiii'  is  Ik'jikI.  liistiiloiriinily  llic  jtlaml  picsrnts  ii  picture  very  liK' 
Ihiit  wliii'li  lias  liiM'ii  (Irsi'iilicil  nlnivc  as  liy piMplasia  ;  lliat  is  to  say,  tlic 
Msiclcs  linvr  a  iJcliriiMii'v  .if  colli. id  malcrial :  tlicir  cpitliclimn  is  coliini- 
liar  aii.l  folded  up  into  tlio  vesicles;  and  tlic  int.isl  ilial  tissue  l.etwein 
llic  vesicles  is  ver\    iiiaikciUy  increased. 

I',\..plitlialiiiii'  J.'.. iter  is  almost  iiniv<'isall\  claiiiied  to  lie  .liie  to  liyper- 
>ecieli.iii  of  llie  llivf.ii.l,  liecallsi':  i  1  tlie  syiii|ito|iis  of  tin'  ilisease  ate  In.t 
iinliki-  tliiise  |ii(idui'e.l  li\  exi'essive  a-lininisti  at  ion  of  thyroid  to  a  normal 
individual;  and  ('J^  tliey  arc  in  "general  ojiposite  in  I'liaia.'ter  to  the  symp- 
toms found  in  cases  where  the  th\r..iil  ..'land  is  atrophied.  The  Mood  of 
a  p.'isiiii  xvith  e\iiphllialiiii."  j,'..iler  i\  hen  injei'tnl  into  mice  in.Teasi's  their 
lesistaiM'i'  t.i  till'  losie  ai'li.iii  III' aeel.iniliile,  \v  liiidi  i^  also  the  .'ase  after 
thyroid  lAtra.-l  has  lieeii  ili.jeete.l.  In  many  eases  of  exophllialliiic  '.'oiter 
partial  reini.val  of  the  '.'laii.l  is  sai'I  lo  ameliorate  the  sympt'iliis.  Other 
clinicians.  h..uever.  slate  that  if  the  patient  is  (.'iveii  jiroper  medical 
treatment    rest,  ami  <Iiet.  ei|nally  I.eiiefieial  results  can  he  . .1. tallied. 

('.■rtain  i n vest ii;a tors,  however,  deny  that  it  has  y.>t  l.e.-n  .•oiicliisively 
demonstrated  that  .'Xophthalmi.'  jroiter  is  due  to  liy|n'rseeretion  of  the  th>- 
roi.l  iMarini'V  It  is  |i.iiiited  out  that,  if  h\  pcrsiM-retifni  w.'r.'  the  I'aiise  of 
the  ilisease,  one  wmilil  expect  that  the  iii.iectioii  into  animals  of  the  Idood 
of  patients  sulTeriii!.'  from  it  \\ould  ))ro<luce  s,\iiiploms  similar  to  those 
following  the  injection  of  thyroid  extract.  The  results  of  such  expori- 
iiieiits,  however,  have  lieeii  oxtreiiiely  confusiiitr  and  very  iiideeisivo,  since 
it  is  dirtli'iilt  to  iei"o._'ni/e  in  lahoratory  animals  many  of  the  cliiiraeteristie 
sym|itoms.  es|  i'  lally  those  atTeeliii".'  the  sl<in  and  eyes  and  the  »reneral 
liodily  nutrition.  Another  dilVicnlty  in  acee|itin^'  the  hypersecretion  liypnth- 
esis  is  Ih.'  I'a.'t  that  an  extra. -t  of  a  '_'lan.l  removed  from  an  exoiihthalinic 
liatieiii  has  no  .lifferent  pli\  si.'K.  ;i.'  a.-ti.m  .ni  a  normal  animal  froiii  nn 
extract  of  ;i  normal  u'land  eontainiii);  the  same  ))ereeiitaire  of  iodine. 
The  evi.lenee  is  l.y  no  means  conclusive  .me  way  or  the  otlier,  ntid  it  may 
well  lie  that  the  oliserxe.I  chanircs  in  the  thyroid  irland  are  not  the  cause 
of  the  symptoms  of  cx.iph'halmic  '.'.liter,  liut  iiii'icly.  like  the  other  symj)- 
toiiis  of  this  .lisea--.'   a  i.'siill  .if  soiii.'  .•uniiili.ni  lo.Mled  elseuher.'. 

The  Relationship  of  the  Thyroid  with  Othet  Endocrine  Organs 

1.  With  the  Generative  Organs. — Evidence  of  an  association  between 
the  female  <_'i'nerative  or'_';ins  and  the  thyroid  is  verv  strong;  thus,  the 


li 


%    II 


''•HI'  'im'' 


■1 ' 


7r>8 


Tin;    INIMii  KIM.    KKCANS.    UK    IHl  Tl.i:ss    (II, ANUS 


tliyrcii(l  lnTdiiics  cnhiiiiccl  at  |)ulnTly,  diiiiiii.'  llir  iiifiiscs,  ;iiiil  diiriipj: 
])roi{iiiiiic\ ,  iiinl  ill  tli\  roi'li  rtniiii/i'il  viiuiiu'  aniiiiJiU  llic  scwial  ulaiiiN 
fail  to  (|c\.      p  iirii|irrl\ , 

■2.  With  the  Adrenal  Glands.      Sc  !■  pau''  74(i.  > 

II.  With  the  Pituitary  Body.     .\rti  r  icmcval  ot'  tin'  ili>ini<l.  ilic  iiitu 

itary    1m incs   -ii-ratly   altcicil   aiiil    I'lilaiunj,   ]iaitic-ulail\    tlic    paix   an 

Irricir.  ill  wliii-li  it  i->  not  unruiiiiiion  to  liinl  lliat  a  iMi-laiii  aiiuniiit  ni' 
\csicl<'>  ccintaiiiiiii;  I'nlloi.l.  imt  iinlil.c  llio^i'  lO'  llic  tliyroiij,  Ih'cuhic  ilrvcl 
opn!.  'I'ITk  (■niliiiil  iiialri-ial.  Ihih  i'\  iT,  ijiics  mil  cnntaiii  iniliin'.  It  is  sai'l 
tliat  this  iiici-caNi'  nf  llir  piliiilai\  al'lfi-  tli\  idicli'domy  does  not  ocriir  i  i' 
tiiyi'oid  cxliacl  ln'  adiiiinisliii  i|.  iiicicascil  artivily  of  tlif  |)aiN  inter 
iiieilia  ol'  tlic  piluitaiy  i-.  al.o  ipiile  plain.  'I'lioe  Tacts  woiilil  at  lii'sl 
.si'_dit    st'eiii  to   indieat.'  tlial    llie  pilui)ar>    .-ind   liie   lli\roiil   can   act    \  ica 

i-ioiisly,  Imt    tliis   is   \ei\    doiililTnl.   I'ni-   ii    has  imiI    1 n    round   that    pitu 

itar.v  extract  lias  aii\  lieiieliciid  el'fi'ct  in  llie  ( ii'al  inent  ..t  yniter  and  iii>  \ 
edema.  \e\  ert  liiless  llie  assiicialion  in  I'linciinu  nf  ilie  I  v\  o  inlands  nnisi 
he  mor-c  or  jess  clo>i'.  m.t  alone  t'nr  tlie  al"ive  reasims.  I>nt  aUo  hecaiise  the\ 
are  linlh  assncialid  to  iiiiicli  ihe  s.mie  th-jree  willi  llii'  s<Aiial  ori;aiis, 
and  liotli  act  on  the  hi'jhei-  linu-lions  i.i  ihe  i!ei'\oiis  svslcni  in  iiiileli  ihi 
sam>'   iiia'ilier. 

■I.  With  the  Thymus  Gland.     'I'Im'   pi  islsie of  Hie  i hymns  in  e\ 

opiillialinic  '_'oiii  r,  as  well  ,is  llie  analoinic  and  emlir.\  ol"iri''  iclalionship 
helweeii  lliyiiiiis  and  lli>  ioi.|.  is  t.aki  n  lo  indie.ite  snmi'  c-!i.se  lelationship 


n  ^ 


m 


1    I 


THE  PARATHYROIDS 

Experimental  Parathyroidectomy 

Mxpcriiiieiila  I     paiai  li,\  roidecloia\     \  ields    results    which     \ary     in    dil' 
I'ereiit   'jroiips  oT  animals,  iindoiilili  dl,\    lieeaiise  of  ihe   fact   lliat   in  soim  . 

siieli   as  the   rat   and   ral.hii,  : ess(,i'\    paiath\roids  ma\    exist.      In    u' n 

eral     ho\\e\ii,    il     has    heeii     foiiml    ihal     if   more    than    two    of    llie    foir 
paratliyroiijs   he    riiiio\ed,    \ei\    ilehnite   and    proiioiiiicctl    iiei\'ms  symp 
toiiis  soon  siipei-veiic'  and   if  .all   foiir  inlands  he  leiiiosed,  a  ipii(d\ly   fat.ai 
result     is    iiie\  italile.      The    most     aciile    svmptuiiis    ale    exliil'ited    hy    the 
earni\ora      They  iii,-i\    not   he  app.iient    for  a  d;i\    oi-  two  after  (he  opera 
lion,  altlioii'jh   durini:  llie   pel  hid   tin-  jinimal   is  in   a   depresseil   stale,   n 
fiisiiMj-  fiMid  and  losiii'j  \\ei',^|i1   r,ipidl\.      The  iniiscles  ale  also  more  i,r  h-s- 
s|  ilV  diiriii'j  this  staue      When  more  ijelinile  snnptoms  appear,  llie\    coii 
sist    of  .-I    markeil    ahnornialil.v    of   muscular   contract  ion,   leadiinr   to   the 
oec-iirienee  of  lihiiii.ir  col  1 1  rael  ii  ii  is.  ,:f  IrcMiois  ami,  later.   In  I'rampdiK' 
ami    clonic    coiil  i  act  imis.      When    spdiiianeoiis    mo\enienls    are    made,    a 


il 


Tin:  TiiVHiiin  asm  rAUATiiYHoin  (jlands 


759 


))(M'iiliiir  sliakiii!:  of  11h>  f(M.t,  like  lliat  iiiadc  l>.v  ii  iioiinal  animal  to  shako 
wator  off  its  i)a(ls,  is  a  cliaractiTist ic  syiiii>t«iiii.  Tiio  slifrliti-st  stimulation 
of  tlio  ])cii|ili<'ral  ncivfs  is  suflli-i.iit  to  iiiducc  one  of  llicsc  attacks,  whicli 

ircnr  Mitli  evor  ini-icasiMu'  fic<|Ui'ii<-.\ ,  \> niinir  at  tln'  saiiio  tinu'  mon- 

l>i'ono\niiT<l  anil  act ipaniiil  liy   otiicr  ilistiiilianccs,  such   as  diarrhea. 

piofusc  salivation,  rapid  pnlsc.  and  ilys| a  lin  the  do<i  hut  not  in  the 

cat).  In  cases  that  are  not  <|uiekly  fatal,  tlie  hair  tends  to  he  shed,  ami 
the  teeth  to  he  improperly  ealcilied  i  in  yoiiiii,'  animals).     Where  a  certain 

amount  of  |)aratliyroid  ti-^iie  lias  1 n  left      for  example,  one  of  the  four 

lohes     the  s\m|  Suns  ma\    not   iippiar  except   under  eouditions  of  si)ecial 

strain    to    the   animal    e( omy,   such    as   pregnancy   or    improper   diet. 

Thus,  in  a  hit(di  fiom  whieli  tliree  of  tlie  four  '.dauds  had  I n  renn)ved, 

no  symi>toms  of  tetany  occurred  until  slie  lie.-ame  lu'cirnant.  Tiider  tin' 
sanu'  c<in<liti..ns  it  has  heeii  foiuid  that  a  diet  of  llesh  is  much  more  ai)t 
to  hrini:  ahont  the  coiidiliou  tlian  nui-  e.f  veu'etahles  or  milk. 

Tetan\-.  as  tin'  ahove  condition  is  called,  may  al-o  hecono"  develoi"''' 
in  man  either  as  the  rcMilt  of  sur-ical  removal  of  the  parathyroids  or 
hecausi^  of  their  imiiroper-  dc\  elopnent,  Tlie  symptoms  in  man  are  very 
similar  to  tliose  ohserved  iu  lahoratnry  aiiiin;iU.  the  oidy  dilTerelice  lieiu'-' 
that  the  muscular  contractions  ;ire  nuu-e  likd.v  to  he  tonic  in  character. 
Certain  symptoms  that  tiuiy  -lev.'loii  .luriui:  preu'iiancy  or  in  the  course 
of  infections  diseases  or  iu  niwhiiru  int'auts  have  also  heen  found  to  he 
associated  with  deu'eneratiou  of  or  hcmoirhai;i'  into  the  paraliiyfud 
(idiopathic  tetany \  and  certain  ohscnri'  iiei\cius  .liseases  in  adidts, 
such  as  iiaralysis  aL'itaiis.  may  i...ssihly  also  \,r  .-issociatcl  with  (duin<res 
in  this  v'land.  Chorea,  epilepsy,  and  eclampsia  have  likewise  heen 
tliouiiht  to  he  associated   with  it. 

The  parathyroid  liland.  hesides  inllueiicinL'  tiu^  nerve  centers,  has  also 
an  itdlnonce  on  iiu'taholisin.  The  symiitoius  pioduced  are:  ( 1 'I  rapid 
emaciation  and  failure  to  u'l'ow  :  i -J '  a  tcmli'iicy  to  the  production  of 
glycosuria,  (d'teii  detect<Hl  hy  tiuiliiii:  that  tlo'  assimilation  limit  for 
caVhiihy, Irate  islnuercl  i  iiairc  (i.".-J  '  ;  and  :! .  in.ist  dctinit.'ly  of  all.  an 
intert'ereuce  with  (•••dciuin  im-talMilisin.  .-is  illustrated  hy  the  failure  of 
the  teeth  and  hones  to  calcit'y  ))roperly.  This  inteifereni'e  with  nrwmal 
metnholism  led  KelloL'u  and  Voeu'lliu'  l.i  ^tudy  the  et'f.'ct  i)rodupe(l  on 
parathyroideclomi/cd  aniiiKils  \>\  tln'  adiniMisl  i  aliou  of  c.ilcunu.  It  was 
found  that  the  symptoms  were  considcrahly  amerKU-alcd.  These  authors 
eonelnded  fiom  their  results  thai  the  r^s.i.tial  cause  of  tetany  is  a 
deficiency  of  calcium  in  the  hlood.  It  is  i>ossih!e  however  that  the  hene- 
ticial  action  of  calcium  salts  in  this  condition  is  that  it  decreases  the 
excitnhility  "f  tl-.e  lui'vous  syste'u.  a.n  a.ciii.n,  whic'i  it  is  known  to 
possess. 


!  I 
J  • 


ii 

i' 

r 

1  =L      1 

!      ':  1 
i  : 

ii^"   Ii 


'J  ■  1- 


I ' 


}■  F 


:;, 

■J     i 


7(50 


Tin:    INIMirKINi;   OHCANS.    (Hi    l>r(Tl.l>s   (il.ANliS 


Wlicii  tlic  ti'lMii.v   is  till'  result  of  a  coinpU'tf  cxlirpaliim  of  all  paratliy 
i(ii(i  tissue,  till'  syiiiptoiiis  can   he  eoiiihated   liy   a   suecessfiil   transplaii 
tation  or  '^fal't  of  paiatliyiciid  tiss\ie  made  fi'oiii  an  animal  of  tlie  same 
spreies.     Indeed,  it  lias  lieen  found  tliat  tlie  siieeess  of  a  i;rafl  of  paratliy 
roid  is  assured  only  when   the  irraft    is  derived  from   tlie  saiiu'  kind  of 
animal  as  that   from  uliieli  tlie  paratliyroid  lias  lieen   removed.     Imi)lan 
tation   into  tlio  sulieulanemis  tissm'  of  a   tetany  patient    of  paratliyroid 
tissue  olitninod  fi'esli  ficmi  tiie  ch'Mdliouse  lias  lieen  performed  willi  hem 
lieial  (luteoiiie. 

\oel  Paton,  Findlay  and  'Watson-  liave  reeeiitly  eoiitrihuted   -rreatl,\ 
to  our  l<no\vled!:e  of  tlie  iiliysiolo-^'ic   iiMtliolo<ry  of  tetania    tliyreopriva 
as  the  ahovi'  eoiidition  is  called.     The  syjni>toms  are  not  due  to  any  eon 
dilioii  alTeetinir  the  niuseles  iliemselves.  sinee  they  disajipear  after  see 
tioii   of  the  nerves.      \or  are   they    |)rimarily   dependent    upon    the   een 
hruni  or  eorehellum.  sinee  ablation  of  neither  aholishes  them.     This  does 
not    ini|)ly  thai   seeoiidaiy   involvement   of  the  liiLiher  eeiiters   never  oe 
eiirs:  on  the  eonti'ary,  the  epih'iitiform  eoiivulsioiis  and  disliirlianees  oi 
eipiilihriimi  sometimes  ohser\ed   imlieate  eerehral    oi'   eerehellar   involvi' 
liieiit,   respectively.      This  leaves  some   i)art    of  the   lower   neuron    relle\ 
ai-cs  as  the  site  of  involvement.     It  is  not  the  atVereiit    neuron,  since  1h' 
tremors  and  .jerkiiijrs  iiersisf  afler  section  of  the  posterior  roots,  leaviir.; 
tlie  all'eivnt    neuron   as  the  .-ilTeeted  structure. 

The  foreproiiiii:  eoni'Iiisioii  led  I'aton  aiiil  his  eo-w  mhi'is  to  eomi>are  iIm 
resimnse  of  muscle  ;iiid  ner\c  to  eh'iM  ric  stiiiiulalion  in  normal  aii>l 
liarathyroideitoiiii/ecl  animals,  .\ltlionudi  there  are  considerahie  varin 
tions  in  the  lesjionses  of  a  normal  .'inimal,  they  are  xcry  delinitely  cn 
au'frerated  in  tetany  when  eitliei-  the  motor  neiirnn  m-  the  niuscde  itseh' 
is  stimulated,  the  exaji'-'eration  in  the  latter  <-ase  hi'iii','  de|)endent  upoi 
alterations  in  the  neinal  striU'lures  iner\e  eiidiie.'s"!  in  the  muscle.  TIm 
iiMM'eased  electiie  eseitahility  i-an  not.  iiowexer.  he  ta!<eii  ;is  a  measui' 
of  the  severity  of  the  eondilion,  for  it  nia.\  he  no  mine  marked  in  eases 
in  W  llich  thei-e  is  ill\  oK  cment  of  the  eerehral  hilllisl'here  i  eausinir  epilep 
tifonii   Ills'   than   in  milder  eases. 

As  to  the  cause  of  the  s.xmptoms.  iininy  possihilities  have  to  he  eon 
siclere<1.  Tn  the  first  ]tlace,  no  direct  relationship  exists  hetween  tin 
thyroid  and  ]iarath,\  roid  in  this  coiineelion.  due  cause  miirhl  he  tli'- 
ahsence  of  some  suhslaiiee  which  checks  the  activity  of  the  iiprvoiis  s.\  s 
lem,  some  chalone  in  Scliiifer's  sense.  That  sneh  is  not  the  c.-ise  is  shown 
aiiioii'_'  olhei'  Ihiiius  h.\  the  fael  lh;il  hleediin.'  ami  tralisfusiiijj:  normal 
saline  immedialel.\  lennives  the  sym|)toms  for  some  lime.  Moreover, 
the  TMclaholie  dislnrhances  t^o  on  \'.  hen  tlie  ner\oiis  symptoms  are  sliu'h! 
Il    liaij    pre\iiin>ly    heeii    llmuuhl    l',\     W.    (J.    .VlaiMliniir'    that,   since   syiep 


ii:  TiiYKoiK   \M'  r\UATiiymiip  (;i..\m>s 


761 


toms  like  tliosc  nf  tftaiiy  can  I'f  in 


ilii.'cd  l.v  <l('ticifnc,v  of  .•alciiini  in  llif 


lioi 


(Iv  anil  tlic  symptoms  of  i)ariitliyn.i<l('ctomy  ri 


•licvi'il  liv  administration 


of  this  cation,  calcinm  (Icfii-iiiicy  is 


tlip  cause  of  tlic  synii)1oms. 


Wli 


ot  dciivin;.'  tliat  these  ions  ma 


liavc  some  relationsliip  to  tlie  syiniiti.ms 


N'oi'l   I'aton  asci 


l-il.es  them    to   iiiln.nialltn,    h:i   (,iHn,'>,}i,ir    I  pajre   ti(l-'>).      Til 


evidence    is 
istered  to  ikumi 


as  follows:     n~i    nimnidiiie  am 


1   iiicthvl    giianidine   adii 


1  nnimals  produce  sviiii>tonis  that  are  iih'iiti.'al  with  those 


following'  iiarathyroideetoiiiy. 


2^    Tl 


here   is  n   mai 


•ked 


increase   In   1h' 


amoiiii 


t    of  tl 


leso   SI 


ihslaii 


in   t 


tomi/.ed   doirs   and    in   the   urii 


om 
tetaiiv.      ( 


he  hlood  and  mine  of  parathyroidcc 
of  children  suffeiiiiir  from  idiopathi< 
ilectoini/ed    ih 


:r.    Ill   certain  eases  the  serum   of  ])arathyrnidectoiiii/ed   <1o<ts 


cts  tijioii  the  muscles  of  the   Iro.^  sir 


iiuai 


■Iv  to  weak  solutions  of  Lriiani- 


dine  and  methyl    irnaiiii 


inc.      i4i   There  is  a   sfrikinv;  simi 


laritv   in   th 


relatixe  amoii 


mis  of  tile   nil  l-ie.'ellolis 


metaholites  in  the  urine  of  jiarathy- 


roii 


lectc 


mizetl  do'^s  aii< 


if  iioriiiai  animals  iii.ie 


I'ted  with  guanidiiip. 


It  is  conelnded  that  the  iiaratliyri 
•liy   prevelitiu','  its  developmel 


di 


they  iiidhahly  exercise  a  re; 


nuiscles. 


ids  control  the  iiietaliolism  of  nuani- 
it  in  undue  amounts.  In  this  wa> 
llie  tone  of  the  skeletal 


ruhitive  action  ujiou 


Since  it  is  similar  with  ro-rard  to  its  charactrrs  and  metabo- 


lism to  the  condition  follow  im_'  uiyroK 


dectomv 


it  is  helieved  that  disease 


,f  tl 


e  paralhyroii 


is  liie  I 


au^c  of  idioi>athic  1etan\ 


The  Relationship  of  the  Parathyroid  with  Other  Endocrine 

Organs 

We  know  very  iitllc  of  the  irlalii'msiiii)  of  the  i.arathyroid  with  other 
eudocriue  organs.  Vincent  and  others  have  stated  that  after  removal 
of  the  thyroid  itself  .•nlar-.iiHUt  of  the  (.arathyrohl  may  occur  with  the 
formation  of  colloid  material  lulu  ecu  the  rows  nf  cells,  hut  tlie  eon- 
clusiou  that  this  rei.reseiits  a  vicarious  function  I.clween  the  tliyroid  ami 
l.arathyroul  '.'lands  is  not  -ciicrally  aceeptcil.  The  sniMMised  relation- 
ships among  the  parathyroid  aii.l  the  pituitary  and  adrenal  glands  are 
also  baseil  upon   nncertain  e\ideiice. 


ntli 


*  II II 


til  I 


I   :        M 


l:l!l.::)l!lii| 


I    i 


-                  i 

! 

t            5          : 
{ 

:i!i'    . 


(  IIAITKU   I-XXXIV 

Tlir,   I'm  ITAI.'Y    KiiDY 

Structural  Relationships 

Situ;il.'(l  lit  til.'  li.-isr  III'  tlic  lir.-iiu  iiiiil  Ivintr  in  tin-  M'lla  tiiirica,  tin 
|iituitiir\  limly  ill  iii.-iii  lines  imt  \M-ii;li  niiwli  im-iv  tliiiii  iuilf  a  irniiii.  |i 
is  ••i.inui-tnl  witli  llic  Inaiii  I'v  a  I'uiiiifl-sliaiH'il  s(ali<.  liif  iiifiinililmluiii 

(Ml  a< uiif  of  a  iiaHuai  .■lift,  wliidi  rims  across  the  filand  in  an  oMiiiur 

l.laiic.  it  is  ,-in  c.-isy  iiuitlrr  to  split  it  into  two  iM.rtions,  an  anterior,  m 
I)ars  irlaiiihilaiis.  ami  a  |iosleriiir,  m-  jiiirs  nervosa.  Tliis  elet't  in  tin 
ease  of  man  is  nsiially  fmnul  to  lie  mmv  or  less  lnokeii  np  into  isolate- 
i->sts  cimtaiiiiiiir  a  eulloiiMike  niatei-ial,  ami  it  represents  tlie  remains  ni 
tlie  (iriuinal  tiilnilai-  st  iiietnie  fmrn  wliieli  tlie  pars  -;lamlnlaris  is  ile 
veloped  :  naiMel,\.  ii  imiieli   uinuiim'  out    from  the  Kueeal  eetoilerm. 

On  jiistnlnui.-  .■xamin.iliiiii  it  will  I'e  I'mnnl  tliat  the  /mrv  <il<iii<liiliin^ 
ronsists  iif  masses  nf  epithelial  eells  with  lar-e  sinus-liUe  hlooil  capil 
jjirirs  iviii-  hel  Willi  them.  These  l.looil  vessels  aro  very  numerous,  s- 
that  in  an  injeeteil  -laml  this  portion  of  the  i.ituitary  stiinds  out  vcr,\ 
prominently.  The  vessels  are  ilerivo.1  frum  aliout  twenty  smalt  art(M'iol<- 
that  eimvertre  ti.wanl  the  pituitary   from  the  cirele  of  Willis,  ami  ontev 

the  (.'laml  l.y  the  infumlilnilu r  stalk  hy  wliieji  the  -.'laml  is  eonnceteil 

with   the  hase  of  the  l.i-ain.     Three  types  nf  eell   ean   lie  ilitTorentiateil 
nDnstainin-r  (ehn.mapliiihe  i   iiml  siraniilar  ( ehromaphill.  of  whieh  latter 
there  are  eells  with  ai-iil-stMininu'  and  others  with  Iiase-stnininir  srramdes. 
the    t'lirmer    heini;    hy    tar    the    nnn-e    numerous    i  Sehiifer ' ."'      In    sonn 
jinimals  sueh  as  the  eat.  the  rells  of  the  jiars  anterior  ;\re  iirranfred  aromni 
the  Itlni.d  sinuses  in   rous  lis  in  a   eulumnar  epithelium.     The  eells  with 
aeid-stainini;  irraniiles  are  said  to  l.eiMime  n.ueh  inereased  in  nuniher  in 
prejrminey  and  aKn  in  the  eiii.nurd  'jliind  nf  ;i.Tniiieu;ily   'see  p;i!,'e  TT".' 
After   thvinideetniiiy    it    has   heeii    uliserved    that    enllniddike    masses    ae 
I'umulale    in    the    pais   ulandiihiiis.   the   eells  smiietiun's  arranu'inj.'  them 
selves  aroinid   these  masses  as  in   the  thymid   uland.     The  enllnid,  h«\\ 
r\  er,  eontains   lln   imline. 

The  jtosterior  part  of  the  '^land.  or  jxirs  mrriisii,  is  eoinposod  almost 
entirely  of  neunejiia.  eeiis.  ;iiid  iihers,  usually  with  some  lixalitie  or 
■jrannlar  mateiial  lyinir  hetweeii  them.  p;iitieulaily    .n  the  nei<_dili(U'ho.Ml 

762 


Till      I'lll   IIAliV    I'.iH'N 


763 


he    IriH-fi 


i.f  till'  iiifiMKliliuliiiii.  into  uliii'li   it    in.iv 

tl„.  artiv.'   lillnriplr  nf  tlu-   '-'laiMl    is   iciuvsilit.'. 


1.      It    is   Ik 


iicvcl  that 


I  l)v  this  iiiiitfri 


The 


i|i|i|y  cil'  til"'  I 


liir-.    l|i'l'Vn>il 


Is  ri'iiitivi'iy 


■ant  \  . 


T'lCtWCCII 


tlic  i>ais  iicr' 


,1  til.'  intiairlaiKlulav  I'lcl't  aliov.-  rctVrri'i 


to  is  a 


lavcr  ot'  (•( 


lis  (lifl't'riny  Ifoin  tliosi 


il   >'i 


thcr  tlic  aiitciior  or 


flic 


postrl'lor 


ilic.     Tliis  la.MT 


>listl 


tulcs  till'   so-i-all('(l   l>'ii:i  iDtif- 


illii.    'i'iit'  cflls  ari'  sdiin 


,  liat  ril«'  lliosi'  of  til.-  liai> 


tliat  tln'.v  air  ills 
\arii'l\,  tliat  i-  to  say 


tiii.-tlv  uraniilar,  tli.'  ■^rainili'^  l"'ii 


rlaiidillaris,  except 
._r  ot'  the   iieiitl'01)hile 


liie\-  stain  w  itii  lieitlicr  lia-K' 


llol-  aeli!  ^lye^ 


\Vi 


•lined   \r-h'l( 


iiitaininir  an  o\i 


)hili Iloid  III 


itei-ial  are  (d'teii   foiiin 


i. 


1         ,1.  ,1   -,,11-1.   tl'-'iK      ''  ,    ■  ifiii-'i^    »:'■'"'' 
lu.     I"    .       I'l.-Aii.^-    li"l"     ■    P    '•■■tl.ii-   ;;'    '■    .■'■""'    '.'^  v,i,lr.,U--     /     lT,r-    Kl,r.,liil.l'i«;    e.    mill"- 


lrv>  ahiindant  than  that  ot'  the 


,ll.iiir>.-,    fi-i'ni    II,   '.Il-s   /■.'■■. ■■o/,';7V. 

Iiftueell   them.      The  hi 1  -upplv    is  IllUell 

p,,.s  .dandiilaiis.  Althoiis-l,  ^^ell  separated  hy  tiie  eleft  lioni  the  pars 
„h„i.hilaiis.  the  pars  inteiiiiedia  is  not  .ell  separated  fro,,,  the  ,.ars 
nervosa,  heeaiise  niaiiv  of  its  eells  extend  for  some  distanee  into  the  lat- 
,„,.  ,„,,„,.,„  ,l,e  nenroulial   hl.ers.     Certain  o,'  the  eelU  in  the  pars  inter- 

, li.i   m:iv    he  seen  in   various  stages  of  eonv-rsioi:   n,to  ^h.hular  hval.m' 

l„„lies  or  a  uiannlar  mass  ot  material  iHa>  app.ar  in  them,  in  either 
..;,se  the  eells  nItimai.K  hieak  dou  n.  settinu  free  the  h>  aline  or  -raiiular 
,„„,,,ial.  Nvhieh  is  believed  to  he  the  orh.dn  o|  the  similar  material  al- 
,,,.„,.,  ,l,,„.,.i!„.,l  ,,s  existin-  l.-tv.ee!,  the  lu'iiroulial  fillers  ot  the  pars 
,„,',.vos;,   and   there.on.   iilliinatel.    llndin-   its   u  a>    hy   the   infimdihulnm 


hi     ii 


f  '  ("1  : ' 


Eft     M 


1'.;. !' 


15l 


ii 


ll 


ill     1 


'       I:  J 


laiu  1*         i 

'  l*  '  '    1 
«   1 '     I 


5|-i 


11:1 


Lib     ; 


ill;: 


704 


Tin;  i.NixH  him:  orcans.  or  di  i  ti.i:ss  (ii,ani>s 


int.)  tlu'  tliii^l  vciitii.-lf  of  tlic  liniiii.  Tliesc  liviiliiif  '-'lolnilfs  arc  •jirnlly 
incrc.iMil  iit'lir  tliv  iiiiil<i-ti>iii\.  it  nIhmIcI  lie  iiitMitiuiicd,  liii;illy,  that  ,i! 
til.'  iiiniiriii  lit'  llir  iiitraulaiiiliihir  clri't  tlic  iiitcniifdiaiy  ami  antoi'ii>i 
liciitiiiiis  III'  llii'  iiitiiilaiy  ciiini'  timcllii'i-,  altlinii-ili  tlif  crlls  nt'  cai-li  can 

icailily  lie  .lisliiiL'iii>lic(l  mi  a uiit   of  tlicir  stainini.'  piopfitics.     Tin- 

I)ar>  '^lamliilaiis  ct  iiilciiMnlia  also  cxtciuls  as  a  tliin  layer  ovor  |)art  oi 
llic  |>ai>  lu■l■vo^a  and  aronncj  tlic  iic-k  of  tlic  ulaii<I  at  tlic  iiifniHlilniliiiii 
TlicM'  iclatioii>lii|»  arc  well  sliowii  in  llie  acconipaiiyiii'-'  diairrain  i  Kii; 
VM\  . 

Functions 

( •oiicniiiip,' llif  ruiiclinii  iif  tlic  pituitary,  it  iiia\   1"'  sai<l  in  ireueral  tlia' 

tlie   anterior   loli,'   Inis   an    iiii|iorlalit    ri'lation>liip   to   llie    nutritive    i 

ijitioii  of  tlie  liody  diiiiin;-  i;ro\\tli,  .■specially  of  llio  skelclal  structure- 
and  tliat  tlie  imstcrior  lolic  in-odnccs  a  \ery  ai'livc  aulaeoid  lia\inv  t<i  d- 
\\itli  tlie  jiliysiolo'ji.'  activity  of  uiistripcd  muscle  (ilicr.  Tlic  pars  inter 
media  seems  to  lie  associated  w  illi  llie  |>ostcrior  lolie  in  llie  production  m 

this   aiitacoid.     Tlic    rumMioii    of   tlicsc    tun   parts   v^ill    ilicrcfoic   I n 

sidcred    toiretlier. 

Function  of  the  Pars  Glandularis.  Tlic  facts  eoncernin;.'  tlie  fuiictioi 
of  tlic  pars  ijlandMlaris  lia\e  liecii  '.dcani'il  larujcly  liy  oliscrviiiK  tlie  ef 
fects  prodiiceil  liy  jiarlial  or  coiiipleic  removal  of  tlic  entire  |)itiiitar,\ 
.jiistilicatioii  for  ascriliinu  to  tlic  rciiio\al  of  llie  anterior,  rather  thai 
the  postcriof.  lohe  tin'  results  that  are  ohtained  lieln;:  fiiniishcd  l>y  control 
e\periiiients.  in  which  l>y  icmosal  of  the  i>os1erior  lohe  a^>lle  siinilai 
effects  are  not  oliseincd. 

roiiiplete  reiiio\.-il  of  the  jiit uit .M r.\  is  almost  iiivai-ialil\  f;itid,  the  con 
ditioii  liciiiy:  calleil  iijiil i(iliiri<iii.  T\Mi  operative  procedures  lia\e  licci 
employed  for  the  removal  of  the  .rhind.  ( >ne  of  these,  elalmrated  hy  CiisliiiiL' 
.iiid  his  pupils.'  ■  consists  in  trcphiiiiiiL'  the  skull  and  c'lcxatiiej;  the  tcmporai 
lohe  of  the  ecichrum  so  jis  to  expose  the  •ilaiid.  The  other,  elahoratcd 
1,\  lliish'\.~  rnnsisis  in  ,ippin;ichiie_'  the  !;l;ind  tlii-oin.di  the  orliilal 
c;i\il.\-.  AllhoiiLdi  IIh  re  is  sumr  daii'.icr  of  injuiy  to  iierxous  tissues  !•> 
the  intracranial  luctlioil,  its  results  are  more  dcpendalilo  since  the  daiid 
is  actii.-dly  e\|)osed  to  view    licfore  liciii'.'  removed. 

^Fost  hypoph.v  sec-iomizeil  animals  die  wilhin  two  or  tlui'c  days,  nnlcs- 
they  are  very  \(iunu-.  This  lonuei-  sui\i\al  of  voiin-r  ;inim;ils  is  asei'ihed 
to  the  iiroenee  of  ;iccessor\   piliiitaiv   nialciial  silu.itcd  in  the  dura  malei 

liiiiii'-'  the  sella  turcii-a.     The  most  c\tciisi\e  oliserv  at  ions  have  liee ad' 

on  do!.'s.  On  the  day  following  the  operation  the  animal  aiiiiejirs  ahoiit 
normal,  hut  it  siailiially  liecomes  less  active,  nd'usinir  food  and  resi)ond- 
ini;sU)\\lv   to  stimulation.     It   ■/ladnally  'jels  weaker  and  weaker;  miiscn 


l'\ 


•iiir  I'lii  n\nv   i'''ii'> 


7«r. 


ar  tn'iiKiis  iiKiv  jiiipi'ar.  Iln'  ns|iir;ili<iii  jii 


i|  iiiilsf  liciMimc  >li>u  ,  tlic  liai-' 


i-li('(l,   tlio   tciiilMiiituif   suliiKinii.il;    :iiiil.    ii>niill.v    u 


illiiii   iiltout    foi-ty- 


hotiis,  <MMua  .l.-vc1..|.s  iiM.l  til.-  aniiii.il  .li<■^  in  tliis  .■nii.liti..ii.     Wli'-n 


tlic  syiiii)tiiiiis  at 


itc    anil    ili'atll    Jims    IkiI     n.-fi 


ir  so   cai-ly,   it 
1  li 


r(\  U    Ciisliiii-  .•Mli.T  lliat   small   pairmiis  .>r  \hv  -Ian. I  liavo  n 

li\ii\   oi' iitlicf  ..ijians  lias  .{.■v.'l.'|i.'.l 


left  iicliin.l  oi'lliat  s.iiii.'  vi.-aii.M\s  a.lni 
1.1  ic|)Iac(-  lliat   i.r  111.'  piliiili 

Wli.'M  ..Illy  a   l>ail    ..r  til.'   i.ilnil.iiy   is   iviiliv.'. 


I  .•itli.r  iiiiint.'iiti.)nall.\ 


.)!•  Ml 


I.Miti.inalh.  111.' ^\  ini.t.iiiis  .-nv  iml   n.'aily  so  aci 


it.',  aii.l  tlio  (•.iiiilili.'ii 


kiiiiwii   as  JiitjHijiil iiiliiiisiii. 

ist  l"a.-ts  .■.iii.'ffiiiiiL;  tlif  riiii.'li.iii 


It    i>   liy   a    stii.Iy   .if  tills  (•()inlili..ii    tliat 
j.il..'  liavc  I'.'n  l.'Min.'.l. 


I  inn  ..r  ill.-  iilitcriin 


Wlicii   til.'  ..p.r: 


iti.iii   is  |icrl'.inii.>l   "11  .\.>niitr  animals, 


llu'v   tail   1.1   'jr.iw 


IH-.-p.-rly ;   tl;.'   milk   l.'.'tli    an.l    lli.'   laiiii'j..   a 

.1.)  ii.it  an\>l..s.>;  111.'  tli.M-.ii.l  an.l   Ui.mmin  .^lan.ls  :ii-.-  .'ii 


ic    r.'taiM.'.l :    tli.'   .■|>ipli\ 


•K   .if  lilt'  siiprai'i'iial   an.l   lli.'   -.AHal    .n-'j-aii- 


lai-i;v'l;  an 
r.-iils   1.1  .l.'V.l.ip. 


'I'l 


il.  tli.iuirli  small,  li.-.-.mi.'s   \.iy    lat    an 


1  mav   tli.Tcr.irc   in.-n'asc   in 


wci^lit.      'j'licro 
results  it 
mitficoiih  liiiviiiii 


(listin.'t    cvi.l.'ii.'i 


..]■    mtiital    .liilliii'ss.       l"r those 


,ii.-lll.le.l    tlial    the   •iii><  ri'if 


hihi    III    llii    jiil liihirii    /( 


!(//( 


1,1   ,1,1    ir:ih    III,    ,},,■,!,, J,, „,  „l    ,,l    III,    all},  lilt   mill   ulhi  r 


stnirtiiria  of  thr  iirmriini 


il.     'I'liat    tlii>  all 


ta.-.ii.l  is  lint  il.Tiv.'.l   from 


tlie  ]>(is1eri.ir 


lolie   is  evi.leiie.'.l    l.y    III.'    fa.-t    lliat    imrtial    iii.iniy 


lolio,  or  in.lood  its  eiitir.'  reiii.iva 


I.   is    11 


if  this 


it    f.ill.iucil  liv  similar  symptoms 


Cl.iser  oxamiiiati.iii   -f  tlu'    m.tali..li.'    finicti.ni    in    liy p,,pli.vs,.et.imi/,e. 


animals  lias  sli.iwn  tliat  ll 
exeliaiiire  of  osyji'ii  an.l  ca 
li/o  carli.ili.Mlrat.'  Iii'.'.niiis 


I'l-e  IS  M  mai-K.'.l  .(.•)ir.'ssiii|i  in  tli.'  respirat.iry 
il..ii,  .li.ixi.le,  1.11.1  tliat  111.'  i'.liility  t.i  iii.'talio- 
li.'iulit.n.Ml;  tliat    is   t.i  say.  tli."  animal   witli- 


iit  developing-  ul\  cisiiria  eaii   t. 


'i-at.'  a    la 


riii'i'  (|iian1  ;t\'   ot   siii. 


the  normal   animal 


This  .'tT.'.-l    .1 


than 
h 


■ijlte.l   ll'it    so  iiill.-ll   \' 


iliohy.liat.'  metali.ilism   may  how 
ith  111.'  fini.-linii  of  til.'  aiilerior  lolie  a 


'uith  that  ..f  111.'  post.'rior.  f.n-,  as  vx  .■  s|n,ll  s.'e  lat.'r,  CushiiM,'  an.l  his 
pupils  hav.'  foiin.l  that  .'Mra.-t  «\'  tli.'  post.'ri.ir  h.lie  has  a  marke.l  I'tT.'et 
on  111.'  assimikitioii   limit   of  .•arlMilix  .iiate. 

Attempts  hav.'  Iweii  mail.'  t.i  -rafl  llie  i.itiiitary,  espeeially  ill.'  aiil.'i'i.ir 
loll...  into  vari..ns  pavK  of  111.'  I....l>.  H  has  li.'.'ii  f.nin.l.  h<i«  .'vr.  that 
within  a  few  .lays  tli.'  .jrafts  alropliy  an.l  .lis;i|ipear  iinl.ss  tli.Te  has 
lieeii  .'.iiiiph'te  n'moval  ..f  tli.'  pituitaiy  itself,  in  «hieh  eas,'  the  ..-raft 
may  remain  for  a  month  ..r  so  an.l  the  .ilherwise  fatal  outeoiii.'  of  liyp..phy- 
soetoiiiy  he  war.l.'.l  .'IT.  S.niiet  im.'s.  wlier.'  lii.'  irraft  has  niiiain.'.!  f..r  a 
lonu'ertimc,  it  is  sahl  that  a  t.iiip.nai'y  inereas..  in  the  -mwlli  .if  tin- 
l.ili.-ei 


limal  has  he.'ii  n 


ili.-e-l. 


Other   oliserv.'is    liaxc    invest 


i.j.-it.'.i    ill.'    elTe.-ls    ill    n.ilinal    anmiais 


iliinums  .iral  a.lministraiioii  of  |>iin 


itar\   si 


ilistali.'e  or  of  siilieiitaii.'.ius 


I  I 


li 


i  I 


liV^ 


KM 


:r 


i^jl'   M'i<!H 


« 


li    ■!  1i 


!i  If  V  n   ' 


ii;t. 


■nil    iMiiMiiiM    iinc;\\-.  di;  hii  Ti  1  >»  lii  wi'-^^ 


iiiji.fliuii  (.r  .Miiicl.     Tlir  .Mlli.T  i.MilK  unv   iii.l.rmil.'  iiii.l  .•..iil'iisin.j. 

|,i',I    I- „ll\    I'.iiiil^fHiil    |{cili.Tls,iii-'    Ik;s   mi ••■.l.^il    ill    iM.hiliin:    I'roiii    111. 

;,„t<.|lnr   loi,..    ii    siilistiii Mil.',!    hlh.liii.   wlii.'h    acrclcnitcs   irn.wlli    in 

vuuiii;  iihiiiiiils  ;ni.l  is  llM.n-lit   \><  linv  n   imssii.l..  viililr  in   liMsl.iiinn  i|,, 
lii'.-iiiin:  |ii'i>('('ss  ill  w  iiiiiiijs. 

Tctli.'lin  is  invli.il.-il.'d  l.\  .iiv  .iti'M-  iVoiii  iiii  iil.-ulM.li,-  .•\lr;ict  ..T  iIh 
.Mrclullv  isiiliitcl  nntrrinr  lol..-.  Il  miitiiins  1.4  \nv  .•ciil  ><(  i.1i(.s|.Ii,mu~ 
im.l  iiitic-cii  ill  liio  |.r,ipnrti,.n  <>\'  U<\w  atoms  l'..r  rvciy  iitnni  of  jthn, 
l)lii)nix.  t"n  .,r  ihc  niliMuc-ii  alniiis  hriii-  |nvsciii  as  amiiN'  i:P>n|>s  aipi 
(iiio  ill  an  iniiiici  jiroup.     Tliv  flVrc-ts  mi  liiowiIi  ..l'  nii.-r  ar.'  in  rvcry  |.ai 

ticular  liki'  lln'sc  ,<<  liir  adininisl  lati'ui  <<\'  antrrior  idl.cs,  I  ccnisist   ii 

f.'tanlatimi  <<(  tlir  liisl  |iniii,,ii  ..|'  tlic  liiinl  'jniv.lli  !•>  (•!.■,■■■'  full.iucl  l.> 
ac.M'lci-alinii  of  llir  latt.r  |,oiti.,n  ol'  lliis  .-yrl.'.  Wii.'ii  fully  m-oun. 
tcllu'lin-fnl    inicc   also  tliffiT    from    normal   animals   in    lirini;    smaller    n 

si/c  Kill  of  greater  wci-lit.  willi  a  ilistinrt  ijiffcir in   ll lulitioM  oi 

llic  coal  Normal  animals  at  foiiilfcn  im.ntlis  of  aye  liavr  •■sliaL'u.\. 
sliirinu  ami  .iiscolorcii  coals. "•  wlicrcas  in  tcllu'lin  f'll  animals  tlicy  liavr 
the  L'loss^  ami  silU\  apiiraraiicc  of  \oniii;  animals.  Dm  in-  liiowtii,  nor 
mal  iuiimals  .ris|,la>  a  -r-atcr  varialiilitv  in  vv.i-lil  tlian  trtlicjin-fci 
animals. 

|-:\traor.ii.:ar.\  effects  lia\c  l.eeii  ol,ser\,'.l  l.v  (lark  to  lie  |.ro,lilee.l 
liy  fee.lini:  la.\  in-  Iniis  \villi  j-iluitary  ulaml.  Tims.  Ky  uivinu:  to  one 
y'ear-oM  hens,  in  addilioii  to  tlieir  iisual  foo.l.  -Jd  millierams  of  lies! 
iiitiiitary  sul-stancc  I'or  foiir  ilaxs,  it  u  as  found  tliat  tln^  a\eraui>  dail\ 
i;iimher"of  e-as  laiil  Us  a  Kateli  of  li.'..".  hens  was  laiscl  from  27:1  .lurnej 
the  four  days  |nccediiiy  tlie  pilnitaiy  feedinu:  to  :1.V2  duriii;.'  Ihi'  foil' 
da>s   of  tlie   administralion,   these   r.'sulls   h.'inLr   ohlaineil    at    a    time    •■' 

\ear  when   the  natural  e-niirodiicti f  the  hens  was  diminisliinj,'.     il 

uas  fiirthcf  oliservcd  that  not  only  is  the  output  of  eu:'-'s  trreatly  iiicieasi  ,1 
as  a   fi'sult   of  the   pituitar>    feedinu-,   Init    likewise   their   fertility,    for   m 

another  experiment  in  which  ;!.".  hciis  \v  ere  kept  alonu'  witli  tw .ckercl- 

of  tlie  same  hreed,  not  only  was  ll utpul  of  euv's  increas.^l  i  from  js  up 

to  :;:;  ,  Imt  the  ferlility  of  the  c'j-s  was  ureallv   cnhai 1. 

Functions  of  the  Posterior  Lobe  or  Pars  Nervosa.     As  already  nun 

tinned,  (•xeisic f  lliis  part   of  the   piluitary  can  he  toleiahly  well   with 

stoiid  liy  the  animal,  so  much  so  indeed  that  from  its  hchavior  al'ter  lli' 
optfatinn  we  can  conclude  little  as  to  the  function  of  the  lohe.  On  ih' 
(itiier  haiul,  extrai-ts  of  the  posterior  lohe  in.jeclcd  into  normal  animal- 
iiro(liice  effects  that  are  \er\    strikinir,  indicatiiiir  that  the  main   function 


inlK^f  icr.Un.-i,-.      Till.,-   t>«lii    ■■.l.^    ■■"'■   f.'-K.    ll"    '■- 
,    ..MH    rcl    l..n 


wliite   rnitf    kt  |  t    ini'Ur    *;inl,!n .  -  ,     ,    ,. 

atlilins    ,ls    m.nMin-ll.:    %,l...,lv    l.,m,.n    -vin    rcl    |..nl!.,n    .Ll-    .hit    l.ioli;      1...    m I,    I...A 

twvnlKim'  .•Ml. I   l«.„tv  iiuht   .l;n-;   .in.l  tlic  lliir.l  .il.nr.t    mn    «,.l,-,   .iltir  «lii.  li   llj.    i.  1...  lu    .1. ,  n  ..-. 
|,i..Brrs^lv.  Iv,    untcl    funli.  r    r.i..ulli   i.a-f,   lictwccn   lliv    lutiitli    .H'l    siMi.ili    «r,  !,,    -ii,..  ..!c!k:    ."' 


Illl     fl n   lI'MtN     IH'I'V 


7fi7 


,f  this  loltc  is  |)n»lii.'ti<.ii  of  all  ailtocniil.     'I'li.'  .aIim.Is  liiivc  iimiv  or  loss  au 


i-pi 


iii'pliriiK'-likc  action.     Siicli  extracts,  rciic 


Icrccl  |)iofcin-l'rcc  ai 


11(1  stcfii 


.(1    nrc  nl.tainalilc  on  tlic  iiiaiUct  niidcr  llic  vMiious  iifiiiios  o|  inHiiiii 


,l)(i|)l 


IVSlll,   (' 


luit  this  is  pioliahlv  a  nil 


tc.    K 
)>aM\ 


I'liiii  tliciii  a  I 


r\  stiilii/.alili'  III 


iit<  rial  has  Keen  olitniiifil. 


i\tiirf  ot'  various  siilistaiiccs. 


In  •)! 


iir  lh 


I'liiiclioiis  of  these  various  extracts,  it  must   lie  iiiiicm 


lirrrd  thill   the  inter 


nipiliaiy  iiart   (jiars   i 
their  iireparatioii. 


nteviiiedin"!    is   iiieliided   witti    the    posterior   lolip    in 


Altlioiuxli  tlie  elTeet  of  iiituitaiv  i 


\tiact   nil  i>Jii!ii  iiiii>irlr  fihi  r   fand  on 


irhindidar  tissue)   ap|)eais,  on 


ly  ei)iuoi>liriiie,   i 


first  siu'ht.  to  lie  very  like  that   )irodiieed 
•xaniiiiatioii   tliat   tlie   two 


has  lieeii    foiiiiil  on  i-loser  i 


llv  act   in  dilTcient  ways.     The  rise  in  lilood  pressui-e  p 


siilistaiiees  rea 

diieed  by   ))itnitar,\'  antacoid   is   lil\( 


cly   to  lip  more  iirolonjieil   than   that 


lirodiiccd  liy  eiiinep 


It  stimulates  iiiereased  cardiac  activity,  hut 


fter  tlie  vairi  have  lieeii  cut  or  sunieient  atropine  admiiiisteicil  to  paia- 
Ivze  thciii,  the  pituitary  autacoid  continues  to  stimulate  the  streiiirth  of 


tiie  lieartheat  without  |>n)diieii 


the  aecpleration  noted   with   epineiihrilie. 


Whereas  epinephrine  has  little  tu"  no  action  on  the  coronary  vessels  i>r 

iitaeoid  usually  pinduees  constriction  of 

if  the  two 


tiiosp  of  the  liiiiv's,  iiiluilaiy  a 


Iioth  types 


if  vessel;   and  on   the  renal   arteries   ihe  actions 


iiitacoii 


are  entirelv  dilTcren 


II 


ii:  elTect,   while  the   |iiluitar; 


it,  I'or  epine]ihrine  has  a  marked  cniistric 
utacoid    produces  dili'tation. 


Aliotiiei'  strik 


ilT 


ni!x  (lilleri'iice  in  i  lie  < 


th 


•xt facts  trnm 


the  two  udands  is  re- 


\ealpd  h\  repeatiiii,'  the  injecti 


ifter  the  elTcct   of  a   previous  on(>  Ihis 


ipletcly   passed    off.      With    eiiinephiinc    the    ori-inal    elfeet    is    rejivo- 


diiccd  :  M  itli  iiiluitiili.  ol 
tioii  is  verv  often  the  ri 


IHcssure,   111 


I  the  other  hand,  the  ciTeet  of  the  s md  iii.icc 

rse  of  that  of  the  llrst;  that  is  to  say,  the  liloi 


itead    of    risiiiir,    mav    tall,    or   tlie   rise    he    very    iiiueli 


iked.     Whetiier  this  effect  of  the  seeoii 


1  dose  is  caused  liv  the  actio 


liiarKed.  \\  liellier  this 
of  an  autacoid  havinu'  a  dalonic  rather  than  a  hoiiiiouic  inilueiice,  or 
whether  it  is  due  to  a  reversed  elTcct  of  the  same  liornione,  it  is  impos- 
>ilile  at  i>resent  to  say.  The  clialonic  etVect  in  any  case  is  much  more 
evanescent  than  the  iioiiiionic,  ami  it  is  not  caused  hy  cliolin,  as  some 
have  sU'.'V'ested.  The  elTect  of  epiii.'iihiinc,  it  will  he  remeinliered,  is 
aholishcd  liy  criiotoxin  and  apocodeine.  These  drims.  on  the  other  hand, 
have  no  inilueiice  on  the  action  of  ])iluitrin.  The  ditVercnce  in  action 
hetweeii  Ihe  two  alltacoids  is  usually  explained  hy  MssUlllinir  that  the 
ppiiiejilirine  acts  on  the  receptor  Mihstaiice  assoei.-ited  in  some  way  with 
teriniiiations  of  the  syiiipathclic  nerve  lilieis  in  involuntary  muscle, 
whereas  pitiutrin  urls  illrnlhi  mi  tin  lurol  iiiilnrii  niiisi'U  jihi  is  tlniiisi  In  s. 
Other  tyiies  of  involuntary  tihei'  are  also  acted  on  iiy  pitintiin.  The 
uterine  contractions  for  example  are  siimulal<'d      Via.  1!'T    :  so  are  those  ot 


m 


i  i 


if 


m 


•  I 


.j 


^il! 


1:1! 


Ttis 


•rill     I  \|Mii  i!IM    (i|(ii\\>.   nit    111  I  I'll  ^^   (.IvShs 


llic  iiiti'stiiH'  I  ill  (•iiiitiii>l  In  till-  iiiliiliiliiii.'  I'lVii-t  111'  ciiiiii'iiliriiH'  I,  iiiiil  ol'  llii' 
lilnil.l.'i-imlcr  iiiiiN.-iiliiliiri'.     I  lilaliiliMii  <,(  tlir  pupil  «(  iIk'  •■M-isnl  U»ii 


1,^,      y.;        I'la.iiig   >lu>\\iMy    llu-   ri<  ti''il   i.t    latiiilrin    (.11    llic-    iittiiiie   coiit)aiii"n^   ami    l.louil    i-ns-tiit 
111    ii    ili.i;,      .M.„li-    ly    I'.arlioin's    tli.  ll.-.l.       iI'h.im    .l.ii  k^.n.  ) 

cvi-  is  pio'lu'-fl      Til"  clT.'ct  ol"  pitiiitiiii  dii  llii'  iiiiisclc  of  tlic  lirinicliinlcs 
is  sliKwii  ill   risr.   l!'^- 

TliP  ijIdiKh  nil  \vlii>-li  111!'  pitiiilriii  lias  tlic  most   pioiidiiiu'od  ai-tioii  an- 


■I  Ml     ITII  IT  \U\    Ili'liN 


;<;!• 


till'  nmninmiy  kI"Ii<I^  ""'I  "'«'  kiiliit-ys.  TIh  hIhI  <>"  ""  *"'»<'/  is  cvi 
.U'lXTil  l)y  tlu'  rtiiinikiililf  iiKTi-asf  in  tlu'  iiriiiuiy  How  f«»llouiiik'  iiijf<'ti<)ii 
of  till-  pituitiiii.  Thin  iliiiifsis  iiiiKlit  of  coiirHc  he  ilue  mti.ly  to  tli<' 
vasodilntntioii  tliiit  wi-  have  s.fii  surh  t-xtfartH  pfoilnci"  a  vas.Mlilatation 
wliich  is  all  tlio  niori-  iiiaikftl  Ii«.vh\isc  the  vesHcls  i-IsowIkti-  in  tin-  l>o<l.\ 
niiiliTKo  I'oiiHtiictioii.  Hut  pituitiiii  i-oiitiiiu«'s  to  cause  iiicri'axr.l  iniiiaiv 
nlitllow  ill  the  alisrli.T  of  iiiiy  .l.'iiii>iislriilil''  \awular  riuiiip' ;  it  aUo  arts 
after  the  ailniiiiistrutioii  of  atiopiiic,  so  that  it  is  coiisi.liMfd  by  most 
ohsorvors  to  act  on  the  oxcntory  ppitholium  of  the  convoluted  tulmlcs 


i  r 


l.k'     i""      Tiaciiig   »ho«um    tli.'    ron.tricliiig    acti<.n    of    piluJlrin    on    the    l,r..n.tii..l. »    ai.'l    il<    .11. .1 
on  blo...l  i.riMiirc   in   a  spinal  .l.ig.      (From  Jacl<s..n.) 

ill  iimch  the  same  way  as  certain  «Uui-ctics,  like  <liuretin.  This  renal 
horiiionic  action  of  pitnitrin  woiihl  ai)i)eai-  to  he  iiiialoirous  uitli  that  ot 
secretin  on  the  cpitheliiini  of  the  paiicr.'as.  Another  reason  fur  heliev- 
iiii;  lh;it  tin  seereloiy  horiiioiie  is  iii(lepcinleiit  of  lliat  pi  ndiiciiiu:  vaso- 
dilatation of  the  renal  vessels  is  the  fact  that  a  repeated  dose  of  pitnitrin. 
althouRh,  as  we  have  seen,  it  usually  has  a  depressor  action  on  the  blood 
vessels,  still  inoduces  a  stinnilatinp:  effect  on  the  excretion  of  urine. 
The  value  of  i)itui1rin  as  a  diuretic  in  clinical  practice  is  now  well 
recognized. 

The  effect  on  milk  sf<rrtii»i  is  best  demonstrated  by  i)laciiitr  ;i  cannula 


III 


;  1^ 


'I 


!l 


y. 
■  ^ 


liiii 


i      I- 


1    ' 

>  i 

\ 

! 

77(1  Till    immhuim   oKiiws,  im  m  (tiikk  t)i.,\S'i>M 

in  111.'  inMiiiiiiiiry  .liiri>  s..  iI.mI  IIu'  milk  miiy  UvvU  llovv  out.  lly  oI.mtv- 
inif  llif  ml.'  ..r  ..UIII..U  .lii.iiiir  llu^  iii.i.<-ti"n  ..f  pitiiiliiii.  it  «ill  lu-  f..iiii>l 
tliat  II  ifiiiaik.il.li'  iiMT.iiNi.  ...Tills  Al'l.r  tliis  in.Tfiiv.l  H<'<M'.'ti..ii  lias 
.•.iis.'.l.   li.uM'x.r.  111.'   iii.i.>-ti..ii   ..f  iii.>n'   piliiitiiii   liiix   ii"   I'mtli.T  flTi-ft. 

in.Ji.iUiii'-'  tliiil  til.'  iiillii.n.'t'  ..I'  ll'f  liiHt  iii.if.'li..ii  iiiiihI  Iiiivc  I n,  not  s" 

nni.'li  I.,  sliiiiiiliit.'  til.'  sc.'ri'ti..ii  i.l'  milk,  iih  I.i  iiwi'l.M'att'  tlif  t.iilll.iw  <>f 

lli.it    «lii.li   |.i'.'\i.iii~l.v   ha.l  1 II  s.'.'ivt.'.l  im.l  lia.l  .•. 'I.'.t.'.l  in  tl..'  alv...li 

iiii.l  .111.  IX.  This  »'lT.<-t  I'NplMins  wliy  tlif  pitiiitaiy  Kaln.'lHu'.ik'ii.'  Nlmiil-l 
Imvf  v.iy  llltl.'  if  any  flT.'.-t  ..ii  tli.'  t..tal  pmiliirlioii  ••I"  milk  ..r  .>ii  tlif 
t..lal  am. .mil   ..I'  I'al   aii.l  ..tli.f  .'..iislitiiciils  c.iiliiiii.'.l   in   it.      lli>t.'l.>i{i<' 

,.\jii„iiiati 'I'  s.'.'ti.iiis  ..I'  a   icNiiiiu  iiiaiiimaiy   irlan.l  ami  .>l'  llu"  sa 

U'laii.l  al't.f  a.liniiiistrati..ii  ..t'  tlif  pitiiitiiii,  lifurs  i.iif  llif  ali..v.'  intfiprf- 
t.'iti'.n  (if  111.'  a.'ti..ii.  Alvf.ili  in  tlif  ifstinir  stalf  will  l>f  r.'iiii'l  larjffly 
.lisl.ii.lf.l  with  milU  aii.l  tlif  fpitlifliiim  tlatt.'iif.l  auaiiist  tlif  l.asal  iiifiii- 

l„i ,  ulifi-.'as  alv.'.-li  fr..iii  tlif  t:laii.l  aftfr  pituitary  aftivity  sln.w  small 

sliiivflf.l-up  alvfoli,  itaiiiiiii.'  liltlf  milk,  ami  willi  fiiitlifliiim  that   is 

w.'ll  mark. '.I  ami  stamls  ..iit  pi'..miii.'iitly  fnmi  ihf  l.asal  m.'ml.iaiif. 

Th.'sf  lafts  takfii  t.iu'fthfi-  in.lifatf  that  iiitiiitrin  stiniulntfs  llif  miis- 
fiilar  (ih.'i-s  (if  Ihf  .lii.-ls  ..f  tlif  mammary  irlamls,  thus  s<|U.'f/.inir  "Ht  |1'"* 
milk  f.iiitaiii.'.l  in  tlifin.  MiiM-n'ir  fil.fis  liavf  li.-fii  <|fsf  ril.f.l  as  fxistiiiK 
L.tuffii  Ihf  l.asal  mfml.niiif  aii.l  fpilh.lial  .-flls,  mii.-h  in  thf  saiiif  way 
as  th.y  .1..  ill  thf  fasf  ..f  thf  swfat  -.'lamls.  At  h'ast  Sfliiiffr  has  Mif- 
(•...■.I.'.l  in  .|fm..iistiiitiiii.'  in  lliis  p..siti..ii  r.Ml-shapf.l  iiiiflfi  nhifh  pr..l>- 
al.ly  l..l..n«  I.,  miis.iiiar  lil.fi's."'  I'.y  thfir  f..iitia.'ti«.ii,  thf  milk  in  th." 
nivi'..li  is  fxpt'll..!  int.'  Mif  .liifts.  It  has  ills..  I.ffii  f..iin<l  that  piliiitriii 
st.miilatfs  thf  s.fr.'ti..ii  <.f  f.'rfl.rnspinal  lliii.l,  an.l  that  this  stimulation 
is  iii.lfpfii.lfiil  of  a  lisf  in  hi... 1.1  prissurf. 

ritiiitiin  has  a  .listim-t  *.//"'  "»  ((irlmli  iiilnih  nn  lahitllsiii.  Aftfr  its 
ill!  iii\  .nous  or  suliiiilaiif.ius  in.ii'.Mi.in,  a  markftl  l..\vfrinj:  in  the  tolfr- 
iiii.-.  \\<v  siiL'ar  Is  i.hsi.fv.'il  I  pM'/.'  (i.'ij).  usually  t.i  such  an  fxt.'iit  that 
..'ly.'..sniia  I.e.-.. Ml. s  fslal.lishf.l.  Ciishiiiu'  an.l  his  pupils  ha\f  foiiflii.l.'.l 
that  Ih.'  p..stfii..r  1..1..'  .•..ntril.iit.s  an  aiita<-..i.l  whi.-h  stimulatfs  Ihf  iitili- 
/„lj f  siiirar  in  thf  li.>.ly.  ( ■.inliiiiiat.>ry  fvi.lfiWf  for  this  view  is  fur- 
nish.'.1  liy  the  .ilisfrvalhin  that  m.'.'hani.-al  stimiilatioii  of  thf  i).>st.'rior 
loll.',  sufh  as  is  pr...lii.'f(l  hy  iiiin.'tnriiii.'  it  with  a  iiffdlf,  is  followed  I>y 
a  t.iiiporary  ulycisiiria,  which  is  sai.l  t.i  hf  as  in'.inouncfd  as  that  fol- 
li.win'.'  iiiiii.liiiv  of  th.'  .lialMti.'  .•.■iit.'r  '  paj.'.'  ti'lM,  pn.\  i.l.'.l  .^'lyc.vr.'n  is 
jvii'Sfiit  ill  Ihf  liv.r.  Thf  pr.Mlu.-ti.>ii  of  this  farliohy.lratf  autaf.ii.l  w.)ul.l 
ajipt-ar  to  lie  iimlcr  thf  font  ml  of  thf  sympathftif  iifrvoiis  system,  for  il 
has  hr.'ii  foim.l  l.y  I'lishinsz  ;m.l  ^-tl.-r-^  l!i;.t  •ctiiiii:!:il!o!!  of  thf  supfri'T 
ffrvifal  traii'/lion,  whifh  has  hffii  known  for  many  vfars  to  he  frf- 
(pu'iitly  foli.iwf.l  hy  >;lyfosiiria,  has  this  f (Tfft  only  provi.lfd  the  postfrior 


Till     IITI   ITVKY    ItohV 


771 


lul,  of  111.-  |.ifiiilii.>  is  ihliKl  Kv.ii  ^*nl•ui.•ill  iiiimipiiliili.ni  "f  llir  pitui- 
tiiry  iiiii\  fNritf  ;i  li,\  p.isni.  li..M  .il"  pifiiilriii,  wlii'-li  wmilil  niT.Miiit  for 
tlic    iflvroMiriii    nit.n    ..lis'  iillir    .xpfiiiiMiitiil    ts.MNi.iii    or    piirtial 

.|.•^trl|i•tioll  ofllir  piluitjii.  .  mil.ir  irritfilioii  niny  l>f  m'I  up  in  ■liM'iiM' 
of  till'  v'liin'l. 

Tlir  L'ly<MiMiri;i  "lii-'li  i-^  ummiII.n  ol.s.rvf.l  nft.'r  partiiii  li.\  popli.vsr.Moniy 

s ,   passes  otr,  to  !..■   follovM'.l  l.y   a    |>.riiiaiii'Mt   .M.ii.lition   of  in.Mcasf.i 

tolciiiiMT  for  siitfar,  I aiisc  iiou   less  pitiiitriii   is  Ixinu'  pro.lu. •(■.!.     It   is 

saiil  lliat  iliuini;  tlic  slau'c  of  inrvcasr.i  toliiaii lialicltH  i-aii  not  In-  pro- 

iliK-nl  .vi'ii  l.y  cNrision  of  tlic  paiu-n-aH.  Tlic  u-lycostiria  pro.lii.T.l  liy 
irritation  of  tin-  posterior  lol.c  is  ae.-oiripaiiie.l  \>y  i\  marked  pol\  iiria  (<iia- 
lietes  iiisipidusi,  uliirli  may  outlast   the  ({l.\  eosuria. 


!      \ 


1.1 


Clinical  Characteristics 

I'.eeause  of  tlieir  importance  from  a  pliv  sioloirie  standpoint,  we  sliall 
now  proeeed  to  review  lirii'tly  soiiu'  of  the  more  important  facts  that  have 
so  far  licen  hrousiht  to  iiulit  hy  clinical  ohscrvalions.  Tin'  patliojoijic 
condition  hn>st  ficiiuently  ol.scrved  alTcctiii!.'  the  pituitary  is  an  adenom- 
atous trrowtli  i«arlicnlaily  located  in  the  anterior  lol.e.  I'.esides  i)ro- 
ducinu'  '-'ciu'ial  symptoms  of  pressui-e,  such  as  diminution  of  the  \isual 
field  and.  iierhaps.  headache,  a  shadow  can  u^nall.v  he  ol>servcd  when  the 
patii'iit    is  examineil   liy   iiu'ans  of  the   \-rays,      (leueral   symptoms,  ccuu- 

n ly  ascrihcd  to  a  hypersecretion  of  the  ant; id  of  the  anieriiu'  lolie  of 

tho  pituitary  jk  riiiliiUnrisiii     lieiriu    sooner    or    later    to    show    tlieiu- 


I 

! 


'I 


\i\ 


li 


'l  I  i\,  1 


772  IHI      INIhkMIM      (lKl.\\>.    iiK    III!  TI.KSS    (il.WhS 

selves.  Tlies*-  sviiiittoins  iiic  .ilmosl  «\;ictl>  npimsitf  in  cliiiructer  to  those 
(ihserveil  in  animals  after  removal  »<  tliis  iw.itioii  ot'  the  Klaud.  Thus, 
the  bones  of  the  extremities  and  of  the  fat-e  heoome  stimulated  to  in- 
eieased  ({rowth,  so  that  if  the  patient  is  youni;,  and  the  epiphyses  there- 
fore not  ossified,  remarkable  elongation  of  the  Ion},'  I'ones  oceurs.  pro- 
dueinf?  the  eoudition  known  as  ;ii<i<iiitism.  On  the  other  hand,  if  the  dis- 
ease does  not  develop  until  aftei'  ossilieation  is  eomplete,  its  elTeets  be- 
eome  most  marked  in  the  bones  of  the   fare,  the  lower  jaw  beeomin).' 


^■' 


li 


I    M 


ii      ! 


■  MAr.  «:        I 


I  ■ 

;     .  i 

;  :  i 

.  1 

l-'iK-    -il'l-      ll.in.l    c.f   a   I'lT-m   all.-clc.l    with    ai  r(.KiiKal> . 

enormously  hypertroi)hie(l  and  the  supraorbital  ridi^es  very  iironiineiit. 
The  lonjr  bones  also  become  eidar-^etl  at  their  extremities,  and  there  may 
be  some  increase  in  lendlh  of  the  vertebral  columi\,  althoujrh  the  statin*' 
does  not  increase  because  of  kyphosis  (curvature  of  the  si)ine).  Thr 
conditifin  is  called  ncrnmrtialii.  Nutritive  disturbances  of  the  skin  and 
hairs  also  become  marked,  causing  the  skin  to  become  dry  and  yellowish, 
and  the  haiis  to  uudcr}:o  almormal  increase  over  the  bmly.  An  earl,\ 
symptom  of  the  (diidition  is  a  failure  of  the  sexual  imWiT  i  Figs.  !!•" 
and  200.) 


Till:    I'lTlITABY    nitDY 


77.T 


After  a  tiiin'  tli.'  disease  \w/n\s  to  aftVct  tlif  pars  iiitonnodia  ct  nervosa, 
and  (listurliancos  in  carbohydratr  iiK-tal.olisni  come  to  he  observed,  eon- 
sistiiisr  usually  in  a  diminished  toh'rauee  accompanied  hy  Rlyeosuria,  in 
the  early  stages  of  the  disease,  folhme.l  hy  increased  tolerance  in  the 
later  sfasies.    The  glycosuria  is  usually  accompanied  hy  marked  polyuria. 
It  should  he  ol>servcd  that  sometimes  tunuu-  of  the  pituitary  has  heeii 
found  to  .'xist  i,osl„iort,m  tliousrh  none  of  the  ahf>vc  symptoms  had  hecn 
rccoi.h'd  durinir  life.    In  these  eas,  s  it  is  prohahle  that  the  .lisea.se  from 
the  start  had  heeii  of  such  a  nature  as  t(.  produce  a  tendency  to  hypo- 
l.itnilarism  rather  than  hyperpituitarism,  for  the  symptoms  are  very  like 
those  ohserved  in  animals  after  partial  or  complete  removal  of  the  tjland. 
If  the  condition  commences  l.efore  a.hdcsccnce,  the  body  fails  to  srow, 
although  the  child  may  continue  to  increase  in  wei<rht  because  of  the 
remarkable  deposition  of  fat  in  the  tissues.    Sexual  developiiK'nt  is  strik- 
iuirly   interfered   with,  aii.l   the  secondary  sexual   characteristies  fail   to 
show  themselves.     In  boys,  for  examide,  the  i>ubic  hairs  fail  to  extend  up 
to  the  umbilicus;  aiul  the  hairs  on  the  chin  do  riot  develop,  whereas  the 
hair  <.f  the  scalp  srrows  jirofusely.    The  bones  remain  of  the  female  type, 
and  a  broad  pel-  is,  rounded  limbs,  small   feet  and  hands  are  often  ob- 
served.    Ill  these  cases  there  is  usually  excessive  tolerance  for  carbohy- 
drates, which  may  expliiin  the  adiiiosity,  sujiar  beinsj  converted  into  fat. 
In    the   liirht    of  the   ex|)erimentid    results,    the    efTeet    on   carbohydrate 
metaboli>M   may  be   explained   as  due   to   involvement    of  the   po.sterior 
lobe.      Mental  development    is  li'tarded,  .ind   psychic   deransrements  are 
sometimes  observed. 

Where  the  hypopituitarism  does  iii>t  develop  until  after  adolescence, 
s.ime  of  th.-  above  symptoms  will  of  course  he  missed,  but  many  will  be 
..bserved.  such  as  dryness  of  the  skin,  loss  of  hair,  and  the  tendency  in 
the  male  to  .-idopt  certain  of  the  female  characteristies,  i)aHicularly  with 
retrard  to  the  jiroxvth  of  hair.  Obesity  and  increased  tolerance  for  sURar 
are  also  evident,  and  ititrnicutation  of  the  skin,  somethinR  like  that  of 
AcMison's  disease,  is  sjii.!  often  to  be  a  i)rominent  feature.  Operative 
interference  in  the  early  staces  in  many  of  these  eases  is  of  undoubted 
benetit,  as  is  shown  by  the  brilliant  work  of  Harvey  Cushint?,  to  which 
the  reader  is  referred  t'or  further  information. 


U       ; 


I      1 


r»       •  i: 


The  Relationship  of  the  PituiUry  Gland  with  Other  Endocrin* 

Organs 

The  relationship  of  the  pituitary  «laiid  with  other  endocrine  organs 
seems  to  be  an  intimate  one. 
1    With  the  Thyroid  and  Parathyroid  Glands.    That  enlarsrement  of 

the  pituitary  occurs  alter  thyroidectomy  in  man  lias  been  known  for  .1 


l      'II, 


If  I    "4     i     i.  .  i!'  i 

'    'I  I  Hi'' 


mj\^ 


774 


Tin:  i:\iMM  KiNi;  iiKd.vN's.  iik  di  «  ti.i;ss  (ii.ands 


I- 

si 

|!  ■ 

h 


m 


1 

1 

i 
1 

1  \ 

1  ; 
i  i 

! 

I  I  'i 


Vj.       ^ 


m 


■It. 


ills  i    '  1 


M:   ! 


(Miiisiclcinlilf  iiiiiiiluT  of  \c.iis,  Tiic  fnlinj;ciiit'iii  iilTi'cts  iiitirc  imrticu- 
liiil.v  tlic  pill's  .•inlciicir.  ;iltlii>in:li  cliiiii'-'cs  ;iic  iilsii  dcscrilit'il  in  llic  pjii's 
iiitciincili;i  el  iicfv  (isii.  Accoiiiiiiiiiyiiiir  tlu'  fiiliiru't'iin'iit  of  tlu'  .•interior 
lolic,  vcsirlcs  fdiitninini;  (•nlldiil-iiiic  iiiiiti'iiiil  nltcii  liccdiiit'  ilex  I'lciiicd  in 
it  liilt  (Mil  iiflci'  tlic  liy|i('iti(ii)liy  lins  iniH-ccdcil  to  il  considtTjililc  <lc- 
Lrii'f,  this  cipliiiid  docs  not  contiiiii  iodiiu-,  inn-  docs  :in  cxtr.ict  liiivc  llic 
s.iiiic  |di\si(i|niric  ciTcct  :is  one  .d'  tlic  liiyroid  vdMiid.  It  ciiii  not  i-c|>t;icc 
tiiyioid  cxtiMct  in  tlic  Ircntnicnt  ol'  )).iticiils  with  u'oitcr  or  niyxcdciiia, 
or  jiiiiclior.itc  the  syinploiiis  ]>rodiiccil  in  ;iniiii;ils  hy  tlic  rcniovMl  of  the 
Ihyiojd  yljiiid.  Deposition  ot  eolJoid-iiUe  ni;iteii;il  in  tlie  piirs  iinterior 
iilso  oeciiis  in  iiiyxedciiiji.  Histologic  idiiinu'es  in  the  jtars  intcriiiedifl  et 
nervosa,  althonudi  h'ss  pronounced  than  in  tlic  ]y,i\y  anterior.  ar<'  iiever- 
tiieless  said  to  lie  pcrt'eetly  distinct  folhiwiiii.'  tliyroidectoiny,  and  to  e(Hi- 
sist  in  an  increase  in  the  hyaline  and  irraiiiilar  masses  wliieli  liave  already 
heeii  deserilicd  as  present  to  a  certain  extent   in  the  liorinal  irhiiid. 

Less  direct  evidence  of  an  association  in  function  lictwcen  the  pituitary 
and  the  thyroid  is  furnished  liy  the  similarity  of  the  effects  i)rodiieod  on 
the  sexual  fiiiictioiis  ami  on  the  treiieial  ilcveloi)iiient  of  yoiiiiir  animals 
li>  the  removal  of  either  uhiml.  In  I'otli  cases  the  animals  fail  to  irrow 
jiropi'i'lv  :  the  sexual  orirans  remain  undeveloped:  and  llie  mental  func- 
tions art'  infantile  in  t>  pe.  In  liypoph\sial  delicieney,  however,  extreme 
adipositx-  is  likely  to  he  more  marUe.l  than  is  the  case  in  cretinism. 

2.  With  the  Sexual  Organs.  That  the  ))ituitary  srland  lias  mudi  1o  do 
with  the  development  cd'  the  sexual  lU'irans  has  already  been  shown.  Fur- 
ther evidence  ot'  a  relationship  lietvvceii  the  sexual  i.daiids  and  the  jiitni- 
taiy  is  furnished  hy  the  follow  iiii:  oliservations.  Alter  eastralioii  oii- 
laiu:emint  occurs  in  thi'  lutuitaiy,  and  on  histoloiric  examination  the 
'_dand  is  found  to  contain  a  lai;:e  niiiiilicr  of  oxyiihile  cells,  particularly 
in  the  liars  anterior.  This  intliicnec  of  the  sexual  ■rlaiids  on  the  i>ituitary 
is  lielieved  to  depend  on  the  interstitial  cells  iiresent  in  them,  for  it  lias 
lieen  found  that  if  the  ovary  m-  testis  is  traiisplaiitecl  into  other  iiarts  id" 
the  l.oily  afii'r  the  castration,  the  chanu'cs  in  the  ))ituitaiy  do  not  occur, 
altlioiiuh.  as  we  shall  sec.  the  1  raiisplaiited  yland  lieeomes  entirely 
atrophied  except  for  the  interstitial  cells.  The  eiilar<remciit  of  the  iiitili- 
lai  V-  diiriiiu  pri"_'iiaiic-y  an  enlaii-'ciiient  which  often  hrinirs  it  1o  two  or 
three  limes  its  iioiiiiMi  wiiu'lit  is  further  ev  ideiiei'  of  its  association 
w  ith  the  ovar.v. 

:'..  With  the  Suprarenals.  .\ssoei;iti m  of  function  is  suusicsted  in  this 
ease  liv  the  fact  that  extracts  of  suprarenal  anil  pituitary  have  very  much 
the  same  etTects  on  involuntar.v  miiscnlar  liher  and  irhindiilar  structures, 
and  it  is  said  that  the  two  extracts  iiiiilually  facilitate  each  other's 
action  ill  this  retrard.     It  should  he  reiiieiiilpered,  however,  that  pitiiitriii 


Pi[ 


ft!' 


Tin:  i-rnrrAKY  t.ody 


775 


an-l  ..piiici.l.rinc  .1..  not   iil-P'-^n-  t"  art   on   cM.Hly   tl,..  same  i.ciiplMTiil 
iiicchaiiisin  (sec  panf  TtiT  i. 

4.  With  the  Isles  of  Langerhans.  sine.,  pituitiin  alT.'cts  .-aiLoliydnilo 
niHalH.lism,  xvliiH.  is  tlidiirlit  to  U-  piimaiily  .•onti-ollcl  hv  tli.-  Isles  ol 
Laniierlians,  it  is  ciaiincd  liy  soiue  ol.sorvers  ti.at  a  .-.■latioMsiiip  also 
,.xist"s  l.ftwc.M.  the  pituitary  aii.l  these  sinietines.  Iiijeetioiis  of  .luo.l.nal 
extraels  are  also  sai.l  to  eause  a  liy|.eis,.eretio..  of  pitwitrin  into  the 
eeielirospinal  Ihliil. 


I     t 


I  i- 


CIIAI'TKH  I, WW 


irJ 


\   '( 


'     ! 


!•    ( 


.'Sll  :    f 


Tin:  I'lNKAl,  CLAM)  AM)  TIIK  (lONADS 

THE  PINEAL  GLAND 

'I'his  pcciilijir  sinic'tuic  lies  Ipctwccii  tli»>  aiitcrior  <'<irpora  ((uadi'ijrem- 
ina.  Miiil  wcisrlis  aliiiiit  1\\ n-tcntlw  of  a  irraiii.  It  is  lai!j;«'st  in  the  early 
veins  ol'  life,  ami  lunlerirDes  retninressive  eliaiiu'es  aftei"  piil)orty.  Micro- 
M'f.pieally  it  emisists  of  epitlielial  cells  ari'anjreil  loosely  in  trabeeula'. 
v'itli  lai'ire  sinns-like  eapillai'ies  lietueen  tin  in;  neurojilia  and  sometimes 
nmsele-filier  eells  are  also  i>i-esent.  Curious  silobules  of  ealearoous  iinit- 
ter  i'l)rain-san(i )  are  also  found,  especially  in  the  jjineal  i:land  of  man. 
Tlie  siland  is  devciopeil  from  an  evafrination  of  tlie  tliird  ventricle,  and 
\t  is  i  .imolo;;ous  ^\  itli  the  so-called  median  eye  of  reptiles. 

The  finirtloiis  of  the  pineal  srland  are  ohscure.  In  cases  where  \\s 
.xlirpatiori  lias  heon  successfully  accom|)iislied  (in  the  fowll,  it  lias  hoeii 
found  tlnit  the  hody  frrouth  is  stimulated  and  the  sexual  charaoteristies 
ileveloped  more  (|uicl<ly.  This  result  would  seem  to  indicate  that  the 
clinical  ohservation  that  tuniois  of  the  pinc.ij  siland  are  associate<l  in 
youiii;  hoys  with  alinorinal  (frowtli  of  the  skeleton  and  with  the  early 
development  of  the  seeoiidar.\  se.xnal  characteristics,  dcjiends  on  the 
t'aet  that  a  condition  of  hypopiucalism  is  produced  l>y  the  frrowlli  of  a 
tumor.  The  iiiniicdiati'  elTect.s  of  the  in.jec-tion  of  extract  of  pineal  ;;land 
are  not  c'laraeteristic,  consisting  merely  of  a  fall  ill  hlood  jiressure,  wliieh 
is,  however,  ohtainahle  when  .-m  extract  of  i>iacticall\  any  eellidar  orfjaii 
is  injecteil.  I'rolon<red  administi  atimi  of  an  extract  to  yrowiiiti  aniinal> 
is  said  to  accelerate  the  growth  and  1o  lirin<r  ahout  a  iirecocious  develop 
ment  of  the  sexual  orpaiis;  hut  this  result  is  somewhat  difficult  to  inter 
prrt,  for,  as  we  have  .just  seen,  similar  ehani;es  o<'ciir  after  experimental 
removal  of  the  eland. 


■^^^^H,     BS ''^  j 

i    i^ 

^Hl  'if 'i  'r 

^^HHii 

■^;l'  n 

1 

THE  GONADS  OR  THE  GENERATIVE  ORGANS 

The  Generative  Glands  of  the  Male 

The  structures  which  are  responsihio  for  the  woll-known  ii)fluencc  of 
the  testicles  on  the  develo|)mei:*  of  the  male  sexual  cliaraeteristies  are 
the   :.o-ealIed    inlei.sl ilial    celts   of    Levdij;,    which    consist    of   polygonal 
shaped  epitlielinl-like  cells,  with  w  cll-niarkecl  nuclei  ,iiid  nucleoli.    Tiijioid 


Tin: 


CINIIAI.    (iI..\M>    AM)    Tin:    (iOSADS 


777 


.'iHi.nl.s.  slMiniii!.'  l.lack   willi  (.sini.'  aci.l,  arc  also  pivs.^iit  m  tlu"  .•ylu- 
l)lasni.    The  .Iftin',.  of  <l.-\ olopnu'iit  of  the  interstitial  .■ells  vanes  m  .lit- 
fereiit  animals,  l.einc  Marked  in  the  eat  and  man  and  ill-marke.l  in  the 
rat  and  rahhit.     In  animals  whieh  show  seasonal  ehanjres  in  sexual  aetiv- 
ity   the  eells  are  most  prominent  between  the  periods  of  sexual  aetivity, 
v.hen  the   seminiferous   .pithelium    is   l.>>  rvident.      They   also   l.eeon.e 
j.rominent    in    <-ases   where    the   semeniferous    epithelium    is    atrophn<d. 
rither  as  a  result  of  disease  .n-  followinir  liiiation  of  the  vas  def.^rens  done 
in  sueh  a  wav  that  the  artery  and  nerves  to  the  testieles  arc  not  ineluded 
in  tlie  lijralure.     When  the  testielc  or  a   portion   of  it    is  urafted  u.to 
another  pa.t  of  the  body,  the  s.^menifeious  ei)ithelium  deffeuerates,  but 
the   int-rstitia!   e.-lls   remain   alive   and   be.M.me   <|uite   prominent.      It    is 
believed  that  the  interstitial  .-ells  are  resi.onsible  f..r  the  i>ro,luetion  of 
MU  autaeoid   that   has  to  do  with   the  .levelopment    of  aeeessory  sexual 
chai'actcristies. 

77m   .inrls  ,.f  .v.slnilinn  an-  not  sitriiili.-ant  in  animals  below  the  v.'rt.'- 
brata      In  all  of  these,  however,  they  are  very  pronounced.     The  cas- 
trated male  frofr  fails  to  show  development  of  the  thumb  pad.  but  this 
.levelopment   immediately  ensues  if  porth.ns  of  testis  from  another  fn.u- 
be  plaee<l  in  the  dorsal  lymph  sae.     In  birds  the  results  are  more  pro- 
nouneed:  in  the  eastrat.-d  mab-  ehi.-k  the  romb,  si-iirs.  wattles,  ete.,  fail  to 
,levelop  but  will  usuailv  do  so  if  some  testis  from  another  bird  is  trans- 
,,1anted   int..  its  tissues.     In  mammals  the  effects  are   most   strikiUR  in 
iMiimals  that   .levelop  marke.l  mab'  .•harart.-risties,  sueh   as  the  >rrowtb 
of  antlers  in  sta-s.    These  fail  to  .leveb.p  pr..perly  and  arc  prematurely 
she<l  after  .>astration.     In  man  also,  as  is  well-known  from  a  study  o( 
,.„nn.-hs,  .-astration  has  a  very  prof..uiid  .'n-eet.    Hair  fails  t..  trn.w  ..n  th.. 
face;  the  larvnx  remains  nn.b'veb.ped;  the  epii.hyses  are  a  loiifT  time  m 
ossifyinjr,  so' that   the  stature  may  beeom.'  '.'leat.  but  at  the  same  time 
the  I'imb  boii.'s  may  b.>  more  d.'ii.-at.'  than  UMial  :  the  sutiir.'s  of  the  skull 
are  slow  in  elosinjr;  an.l  the  wh.ile  arehiteeture  ..f  a  castrated  male  comes 
t.i  be  very  like  that  of  the  female.     Conlirmntory  evidence  of  the  influ- 
eiiee  of  the  testicles  on  the  d.vel..pment  of  secondary  sexual  ch  racter- 
istics  is  afforded  by  the  obs.rv.ition  that  malijinant  tnm.>rs  of  the  testes 
ill  boys  iirc  associated  uitli  the  |.r.niatuiv  development  of  the  seeondar.v 
sexual  .•haracteristi.'s,  and  that   these  may  re.'cde  after  the  removal  .)f 
the  tumor. 

As  a  f'-uit  of  ci>s;  ration,  iiitercstinjr  chaii(res  have  also  been  ob.served 
in  other  ductless  jilaiids.  Thus,  the  suprarenal  cortex  and  the  thymus 
become  enlarsie.l,  whereas  the  thyroid  and  pituitary  become  atrophied. 
The  m.'iabolic  fwiicfions  also  In  VMnie  tnrdy.  as  is  .vid.'uced  by  a  tenden-'v 
to  the  dep.isition  of  fat 


^1  '    '  |i- 


!    '  ! 


H"t:     ill 


778 


Tin:  iNiMnijiNi;  oikians,  dk  hi  <  ti.dss  (jlanks 


1  Vi' 


I  jilljil 


Wlifii  the  (■jiNtiatiiiri  is  pcifoinicil  tm  nil  iidull  man,  tlio  !il)ovo  cliaiiKcs 
ill  the  sexual  cliai-nctfristics  arc  of  ciMirsc  imt  so  cvidt'ut,  nltliiMiirli  the 
jnostaft',  etc.,  atriipliv.  The  cITcct  (iii  the  iiu'lalxilic  functions  is,  how- 
ever, very  iiiarkeil,  tiiere  lieiiij;  a  strikiiitr  feiidencv  to  iiieieasid  foriiia- 
tioii  of  f.it.  It  is  iiiterestiii't  tliat  aeeoiiipanv  iii«  tins  tliero  should  usually 
occii!'  a  loweriiij;  of  the  assiinilation  liiiiit  for  eailiohydi'ate,  so  that  irlyeo- 
siiria  is  very  readily  induced.  We  can  not  assume,  therefore,  as  ("iish- 
iiii.'  has  done  in  the  case  of  hypopituitarism,  that  the  fat  dejiosilion  is 
attendant  ii]ion  an  improper  coiiilnistion  of  carhohydrate, 

'I'licse  reiiiiirkahle  elTects  of  castration  have  naturally  prompted  oh- 
ser\('rs  to  study  the  inllueiice  of  injection  of  testicular  exti'act  on  the 
dcvcIoi>nient  of  sexual  <'liaractcristics  in  dilTerent  animals,  hut  the  re- 
sults have  in  general  heeii  considt'red  to  lie  neifative  in  character. 


■;> 


I  iU 


m 


■  !   1 


iCI 


The  Female  Oenerative  Organs 

Tt  is  well  known  that,  hcsidcs  their  function  in  producing  ova,  the 
ovaries  also  produce  autacoids  that  liaM-  to  do  not  only  with  the  fixa- 
tion of  the  ciiihryo  In  iilmi.  hut  also  with  the  chan^'cs  that  occur  during' 
l>re!riiaiii-y  in  the  maternal  oiiriinism.  It  is  however  at  jiresent  uncertain 
iis  to  where  these  autacoids  arc  i)rodiiccd  in  the  ovary.  The  two  most 
likely  soiiices  are  the  stroniii  cells  and  llie  corpus  luteuni.  In  the  stroma 
of  the  o\ary  of  certain  animals,  <:roups  of  cells  have  heen  deserihed 
lia\iiii,'  a  dilTcrent  appearance  from  those  of  ordinary  stroma  cells. 
Tliey  have  liecii  called  //"  liilirstlliiil  cilln  of  the  ovary,  and  are  helieved 
to  he  analoiroiis  with  the  similar  stru<-tiires  found  in  the  testicle.  It  is 
Iiossilile,  however,  that  these  interstitial  cells  are  iiothiii<;  more  than 
cells  derived  from  i)re\  ious  cori)ora  lutcii.  The  latter  are  formed  hy 
]n-oliferation  of  the  follicular  eiiitheliiiiii  which  remains  after  extrusion 
of  the  ovum,  and  hy  the  injrrow  inn  into  the  follicle  of  the  so-calle<l  fhecii 
cells  iind  hlood  vessels.  The  fully  developed  c(M'1)us  luteum  in  most 
animals  consists  of  cells  arrani.'ed  in  trahecuhe  converjring  toward  the 
sc.'ir  which  formed  at  the  jilace  where  the  follicle  hail  hurst.  The  luteal 
cells,  as  they  are  called,  are  characterized  hy  coiitainiiii;  considerahle 
(piantilies  of  lipoid  material. 

That  the  ovary  prixluccs  some  ;iutacoid  is  evidenced  hy  hotli  clinicjil 
and  ex])erimental  ohscrvations.  Thus,  if  hoth  ovaries  are  roimived  in  a 
yoiinir  animal  (oiiphorectomy  or  spayin<r),  it  is  well  known  that  not 
only  does  the  uterus  fail  to  develop  properly,  hut  the  external  ehanires 
characteristic  of  pulicrty  in  the  female  fail  to  materialize,  although  act- 
ually the  sreiieral  elTects  are  not  so  pronounced  as  they  are  in  the  male 
after  eastralioii.  Menstruation  does  not  set  in;  the  mammary  <r|ands  fail 
to  develop:  iiml  there  is  a  t<'ndeiicy  for  the  hair  to  grow  as  in  the  male. 


tin;    riNK.M.   <iI,ANl)    ANI>    Till.    (iONAHS 


779 


\Vb,.n  tlic  n|M ..Mlioii  is  pcrfoniicl  in  a.lult  lit'.'.  Hm'  clianircs  arc  not  very 
i.n.Moimc.Ml.  ox.M.pt  that  inciistnintion  ceases  an.l  the  iitenis  aii.l  mam- 
niary  ulan.ls  atn.j.hv.  :MetalM.lisni  als..  heonies  alternl,  eaus.nff  a 
te.i.i.ney  I"  the  .lepositioi,  of  fat,  and  in  the  ease  of  the  human  animal  at 
h'ast,  there  is  rre.|ii<'nlly  evi.leiiee  of  mental  disturlmiiee. 

Attempts  to  ae.inire  more  .lellnite   information  ivsrardini;  the  physio- 
lofiie  elT.^ets  of  the  ovarian  autaeoi.l  have  r.eently  heen  ma.le  l.y  SehiitVr 

;iml  lla-aki.'"   Kstr;irts  w.'re  prepiMv.l  fr tiir  eorpils  luteuin  or  Uraalian 

'f„lli,.|es"or  fnm.  the  hilnm  ovariae.  and  ol.servath.ns  were  ma.le  on  the 
HTe.t  prodneed  on  th.-  hehavior  of  the  ehief  for.ns  of  nnst,,;ited  n.usele 
I,v  a.Min-  the  estraets  to  isolated  prei)arations  of  uterus  or  intestine 
or  l,v  inje.-tint;  the  estraets  into  nnimals.  Appli.d  to  the  isolated  prepa- 
rations extract  of  foliieiilar  tissue  or  of  liquor  follieuli  was  found  to 
increase  the  force  and  rate  of  the  rhythmic  contractions  of  the  uterus  as 
well  as  its  tone,  uhcr.'as  inhil.it ion  was  produced  when  extract  of  the 
hilum  was  used.  Kxtract  of  corpus  luteum.  when  injecfd  into  the 
veins  was  found  to  cause  the  ut.'ius  to  increase  its  contraction  or  it 
,,nicscent  to  hcsiiii  .•oulraclinu'.  It  was  further  noted  that  extracts  of 
hilum  .•aiised  a  fall  in  arterial  Mood  pressure,  whereas  those  ..f  corpus 
luteum  had  little  or  no  eflVct.     It  would  appear  from  these  ol.scrvations 

that  the  extracts itain  two  dilTerent  antacids,  one  havinit  a  hornuMiie 

anil  the  other  a  chalonic  action  on   plain  muscular  filter. 

Kxtract  of  corpus  luteum  when  intravenously  injected  also  stimulates 
the  outpourin-;  of  the  milU  from  the  mammary  glands,  although  not  so 
marke.llv  so  as  extract  of  pituitary  u'lan.l.     This  pituitary-like  action  is 
not  ol.tained  with  extracts  of  ovary  that  do  not  contain  eorp<'ni  lutea. 
Itesidcs  l.eini:  concerned  in  the  outpourinir  of  milk,  corpus  luteum  has 
als..  I.ceii  slM.wn  to  l.e  irtate.l  in  some  uay  t..  the  .levcloiu.icnt  of  th. 
mammarv  jilan.I  .lurinc  i.rcirnancy.     Th.'se  srlaii.ls  l.ec.me  dcveh.ped  in 
y..iinfr   viiTin   rahhits   after   th.'   .•..ntinuous   administrati..ii    for   a   month 
or  s."..f  ."xtra.-t   ..f  .-..rpus  lul.-um.  an.l  th.-y   als..  .l.-veh-p  in   unimprc,'- 
nated  animals  v  h.Mi  the  .-..ritus  luteum  is  made  to  .level. .))  l.y  artificial 
means  sn.-li    .s  punctnrin-  th.-  (Iraalian   f..lliele.     Furthermore,  destruc- 
tion of  the  .•..r|...ra  lutea  in  a  i.rei;naiit   rai.l.it   arrests  .levelopment   of 
the  mammary  srlan.ls.     Th.'  .'..ri.us  luteum  has  also  an   important   fuiic- 
lion  in  .■..nMe.-ti..n   with  the  forinati..i4  ..f  the  uterine  deci.lua  and  the 
lixath.n  ..f  the  einl.ry...    Thus,  after  .lestru.-ti..ii  ..f  the  c.rpus  luteum  at 
an  early   i.eriod   in   pr.'.^'iian.-y,  th.'  emi.ry..   fails  t.i  l.e.M.me  a.lhcrent   to 
the  uterus. 


i 


I      :1 


*l  '•!! 


f  -^^ 


H  'i 


i  -' '.;' 


i|-. 


(hll  1111.    IMHICHINI     tiH(iANS.    OK    I>I(TI.1>S   (il.ANOS 

I>t  (  TI.KSS  (il.ANDS   KKI'KIfKM  i:s» 

(Monojiraphs'* 

■i-ViiicTiif.  Swiilr:      |nlcrn;il  Si  .rations  .■iii.l  tin'  l>i!.tl..*  (iliin.N,  101.  .VtiimM.  I,.ini 
•:'lli>-ill:      Till'  InliTiiiil  Stvii'l.itv  «)im;iiis.  Will.  \V..M  I  \   I'"..   I'.il:;. 
■'■'ScliiitVr,  Sir  i;.  A.:      Tin'   Kihlnriiii,.  Oi;;:iri....  I.i.iiyin:ii.^.  li.ivii   \   <'.,..  N,  «    \>.i 
I-oii.|(in.  ]•.>]!'. 


•  h.u. 
I'rk  :iii 


lOriiiirial  l*ii|HM's) 

••'iKiilk,  M.  y...  »iu\  MjuK.mI,  .1.  .1.   i;.:     .\in.  .Iciir.  I'I.vmi.I.,  lUlH.  xl,  L'l. 
'■-Ki'liii.  (»..  Cmiiihjii,  W.  1'...  iiTi.l  l>iMii,.  \V.;     ,1.1111.  liii.l.'  Cliiii...  l:ii:!,  Mii.  1(7. 
•■'I'aiiiKiii,  W.  II.,  mill  (irii.v,  II.:     Am.  ,li.iir.  I'l.v.l,.!..  lull,  \xxi\.  l'.;:';  hK,,  «ii1i  M,.„ 

ilciiliiiil,  \V.  I..:     II, ill..  -i:i  mill  L'.-.l. 
'■••lliirliiiaii,  T.  II..  nil, I  hiIhts:     Am.  .li.iir.  I'Iivm,,!.,  1;i|.-,.  \xx\iii,  I:;.!;  il,i.l..  l!iir.  xliii, 

•  ill  ;   iM.I..  xliv.  .I.",.);  il,i,l.,   llHs,  xlv. 
'■  lIiiskiiiH,  K.  <i.:      Am.  .I,,iii.  I'IivnIlI.,  I'.H:;,  x\i\,  M',.:-.  .|,,iir    I'liiirm.  iiii,|   \.sy.  ■nii'iii|i.. 

I'.'ll.  iii.  !•:::  .\iii.  .I,,iii.  I'liv>i,i|.,  im,-..  x\x\ii.  171:  il.i.l.,  li'lii.  \li.  ."il.;. 
"'■■Slowiirl,  (i.  X.,  :iii.l  I!„-(,ir.  .1.  .M.;     .r„iir.  I.iil,.  tiii,l  Clin.  .Mi.l.,  I'.Us,  iii,  I'mi.     Sor  full 

l,ilili,i:;r;i|'liv  l.v  l{(ii;ii(r  in  tln«  ].ii|>,t. 

fiM'.iiii.tt,  T.  |{.:    J,  111.  riiM-iiii.,  i;mi.',  xliv  :;7i. 

'■S><fc'Wiirl,  (i.  X.:     .I.iiir.  Kxp.  M,',|..  liill.  xjv.  ;'.77:  ili.l..  It'll.',  xv.  HK;  il,i.l..  xvi.  501' 
lii'Stc'W.irt,  <*.  X.,  l{„y..ir,  .1.  M.,  .-iml  (iil,s,.n:     J"Ur.   I'liHini.  jin.l  Kxiut.  Tlicnip.,  l!»lf>, 

viii,  L'(i.">. 
■ 'Mclt/cr.  S.  .1.:      |),'i;t..,l,.  mr.l.  W.hnsilu.,   I'.hih,  xiii. 
:isti'«iiit.  «i.  N.:     .Iiiiir.  Kxpcr.  Mi.l.,  I'.'ll.'.  w.  ."il7. 
■-■(■.•iiiniin,  W.   I!..  I't  ill.:      .\m.  .Imir.   I'livsinl..   r.'ll,  \\\iii.  ill;   il.i.l.,  I:,l  I.  \x\iii.  ;;.*iii: 

iilsii  l'...ililv  ('li.'Miuii  ill    lliiiu,'!.   I'l'iir,  ;iii,|   Hiiu,',  .\|i|,I.t..n,   lHI.'i. 
;  ■•('iiiiiiiiii.  W.  I'.,'  iiilI  r;itl|.ll.  .M.K.-ii:      .\iii.  .I,.ni.   I'li\si',l..   I'.'lf,,  xli.  71. 
••Mn.li'.Hl,  ,1.  .1.  I{..  :,ii,|   |',.:ir,i',  l{.  C:      .\ii:.  .Ii.ui.   I'li'.\.*i,.l..   I'.'ll.'.  \\i\,   ll!>. 
■■■Xliiiiiu',   |i. :      l'i'i>.i.nil  r.iiiiiiiuiii.iiti.'ii. 
■'■Miirini'.  I).:     ,li,iir.  I:\|.it.  Mi',1.,  I'.ill,  xl\,  S'.t. 
■■Mnrin,'.  ll.,  im.l   I.i'iiliiirt,  ('.   II.:     .Ii.ur.   Kxprr.  Mv<\..  Il'ln,  \ii.  ::il:   il.i.l..    p.'ll,   \iii, 

■l."i.";   .-lUi,   Hull.  .I'lliiw   H'lpkiiis   lli,sp..    Illin,    xxi,   Ji.'. 
•"Xriirinc.  1)..  iiii.l    Kim'.:ill.  (1.  I'.:     .1,  ur.  I.jil,.  :in,|  Clin.   Mi,l..   I!il7.  iii,    11. 
;:'Krn.l;ill.  K.I'.:      Iii.>t.,ii  .M.  ,1.  iiml  Stui;.  .I..111..  I'.'Ii',.  1  7.',,  .",.">  7  :  i.U"  I'm,'.  .\iii.  rii>.<i..l, 

Sii,'..  .\ni.  .Iiiiir.   I'livsii.l  .   r.i|v.  sliv. 
S"l'at'iii,    X(„l    nil. I    Kinlnv:      (Jmiit.    .L.i.r.    lAp.    I'livsi..l..    I'.'K.    \.    l.'"".      I'nt,.ii.    .Vi.cl, 

Tinlnv  nml   Wiitw.n'.  A.:      Il.i.l..  L'- ..  Jl -.  ..l  .'■.'iin.l  :;77. 
'iMin'Callnni.   W.   (i..  I't.'.:     .Tiiiir.    Kxp,!.    .\l,.l.,    r.l'i'.l.   xi,    ll'>;    il'iil..    I'.'l:'..    wiii.  HI''.: 

.Ii'iir.    riiiirm.   nml    KxpiT.   Tliri;ip.,    I'.'ll,   ii,    tl.'l. 
'-Cusliiiii;.   IliiiM'v:     Tlii'  ritiiiiiin    1'   .Iv  :iii.l    \u  Pis,, 1, 1,. is.  .1.    I',.   I.ippiii.-..it  (',...   Il'li; 
'-HiTsIi'v.  v.:      Itrit.  Mi'.l.  .I..ui-,.  1^^.",.  i.  111. 
'■iK<il.''its,,n,   Hi.-iilsl'i.r.l,  nn.l  |{:i\ ,   I,.   ,\.:      .I..iir.  Hinl.  I'lifiii..   Il'l'',,   xxiv.  :;17,  "Ii.':,  .'S.".. 

;'.!i7,    Il'l'. 
"■"•Chiik,  I,.  X  :      ,T..iif.  Hi..!.  Clniii..  H'l.-,.  xxii,   l'<,j. 


u.t'i   lliat    li.t    .!;.ta!...!i..!ti 


III 


4 

J 


PART.  IX 
IMF.  (  KNTKAK  NKKVOUS  SYS'IKM 


CllAl'TKU  I.XXW  1 
'UK   FAOI.ITK'N-   «>I'  TIIK   NKISVorS  SYSTKAI 


I  Is  \ 


5  . 


Tl 

tcrs. 


IC    IICl'\<lllS 


siclll 


if  tlic  liiiiluT  aniiuMls  i-c.iisis 


ts  of  llic  luTVc  e<'ii- 


iiin 


1    Ih 


ilh    their    vnrinUN    intiTi' 


iiiufctiti'j:   tracts. 


Tl. 


nerve  traet  and  ci  iitei- 


iited  iiiaiiilv  in  tlie  si.inal  <''ir< 


1  ami  liraiii 


>iiil>lex  stnii' 


xvl.ere,  l.v    their   iMterhieeiMei:t,   tlie.v    f..rn.   an    extnn.el 

„„.e.     The  exae,   position  of  ,1 nters  an.l  tlw  eourse  an.l  -'"'-;"'- 

nf  the  traets  with  the  e..nters  are  ,.rohhn.s  whud,,  un.ler  the  tlth  ol 
„.>„n.lo.y.  have  -lurin.  reeent  >ears  heen  ..ntrih-rte.l  to  '""-  I-'  -- 
l„.lv  hv  the  anaton.ist  an.l  the  ,,atholo.^Mst.  The  n.torn.atu m  ht.s 
„„„•,,,,,;,  t,„s  „s  the  possible  traet  or  traets  of  nerve  til.ers  through  M 
U,o  various  centers  n.a.v  ..,>n,n.uniea.e  either  vvith  one  at.other  or  u.th 
,1,,.  stn.etnres  outsi.le  the  central  nervons  s.v>ten,  upot.  ^^hu.h  they 
,..t  Since  each  of  these  center,  may.  houever.  he  playe.l  upon  l.y  n,- 
flnonees  cnninv  fn.n.  .litTerent  regions  of  the  ho.ly.  it  is  evident  that  there 
„,ust  rcnnin,  as  an  eMually  in.portant  aspect  of  the  suhject.  the  n.vest.- 
.,„ion  of  the  tneans  l.v  uhid,  the  vari.ms  availahle  c<.nters  an.l  tracts  are 
hrouKht   into  eonnnunieation  atnl   a.Mion  at    the   pr,.pn-  tnne.      In  other 

wonls.  we  tnust  inves,ij:ate  //„  f t>n,n,l  ns.s  ..I  lln   „r,nlahh    ,,nth. 

\Vc  mav  rompnfp  the  central  nervous  sysletn  with  a  telephone  systom. 
the  exchansres   representing'  the   nerve   centers,   and    the   wtros  the   nery- 
trunks.     Anv   ineon.in.^  wire   .nay   he   con.ic-ted   l.y   the   operato,-  w-.t 
,,nv  out.'oinsi  wire,  hut  a  knowh.du-e  of  how  each  wiie  runs  .loos  not  tell 

us'undcr  what litions  the  various  ^^i.■es  will  he  ..onnect-d  lor  irans- 

nnssion  of  messa,'es.     It  is  the  sa.uc  with  the  nerv.us  sy>t.-n. :  the  n.M.rolo 
Kist  can  tell  us  how  the  tract,  and  centers  run.  l...t   t.ot   the  cond.  nu.s 
under  which  they  nu.y  aC  t,.,e.her.    Tins  i,  is  the  duty  of  the  phys.olo.wt 

to  ascertain. 

Sine«  It  is  the  degree  of  development  of  the  central  nervous  system 
whieh  determines  an  animaPs  position  in  the  ovolut.onary  scale,  mud. 
informati.m  coneerniun  the  r.^lativo  in.portat.ce  of  the  various  parts  ol 

7S1 


I 


i  , 


■■ 


■  I 


782 


TIM.    CINTinl.    MKMil  S    sYsTKM 


Ht  ■!  ^  i 


1  ,ii':i' 


i/  4  If    . 


\  1?  i 


i} 

.. 

i 

1 

hi 

lA, 

..mL\ 

it  i-nii  lie  iiIcMhi'il  fioiii  a  >iir\cy  "f  llic  <-i)iiililiiiiis  iiihIit  \vlii<')i  tin- 
IH'I'Viills  sVNtciil  liiJikt'H  its  ll|'.|ii'iillllH'('  ill  the  lo\,('st  tnniis  nf  niiiiniil 
lil'i'.  Ill  tlic  ciiM'  lit'  iitiii'i'lliiliir  iii'LMiiiMiiN.  Mii'ii  as  till-  iiinihii,  tlii'  ap 
jilicatidii  >>f  a  stiimiliis  in  tlic  sni'fai'o  i'aiisi'«<  a  iri(i\  inoiit,  Itcpniiso  tlii' 
prntiiplasni  of  tlic  I'luaiiiMii  ikisscsscs,  aniDii'j  its  otlicr  piii|ii'rtit's,  tli"sc 

iif  I'xi'italiilit.v,  I'liiKJiii'tivity  ami  itiartility.     In   tlic  case  i)f  iiiiilticcl 

Iiilar  orL'aMiMns.   on   llic  ntlicf   liainl.   sum lis  arc   set    aside   aiiil   spc 

cializcd  for  tlir  assiinilation  of  food,  others  fur  inovcincnt,  others  !■• 
receive  Ntiiiinli  from  tlic  outside,  ami  >  et  otliers  to  compose  the  touirli'  i- 
tissues  which  jirotcct  (he  surface  of  the  animal  from  injury.  This  loca- 
tion of  spccilic  fnniMion  in  spcci;ili/cd  ".'I'onps  of  cells  mak<'s  it  iieccssar.^'. 
for  the  vclfarc  of  the  or<.'anisiii  as  a  whole,  that  some  means  of  com 
mnnicatioii  should  he  luovidcil  hetwecn  the  distant  jiarts  of  the  animal, 
for  otherwise  tin'  cells  which  are  occnpjcd  in  Mlisorhing  foixl  would  I'c 
linnl)lp  to  move  away  or  he  protected  from  harm  when  some  destructive 
ancncy  ai>proa(di(  d  them,  and  indeed  the  niovinj;  (miisclcl  cells  could 
never  know  when  the  welfare  of  the  ort'anism  as  a  whole  (h'liianded  that 
they  should  heconic  active. 

It  is  proliahic  that,  in  soiiic  of  the  lower  on^anisms.  the  tiicssajrcs  trans 
mitted  from  one  irroiii*  of  cells  to  the  others  are  carried  liy  chemical 
suhstaiiccs  present  in  the  circiilatiiii,'  tliiiil  hormones,  as  they  jirp  called 
fpaiic  72!').  h'or  the  ipiicU  aclaplalion  that  is  necessary  in  the  strusfLde 
for  existence,  however,  such  hiu'iiiones  arc  usually  too  slow  in  liriiinimr 
ahoilt  the  response,  and  very  early  in  the  evolntioiiar.v  scale  we  find  that  cer 
t.'lin   cells  heeoiiie   ditTelelit  latcil    lor  this  sper-ial    ]>Urpose.      The   eclls  thus 

s|)eeiali/ed  constitute  the  nervous  system,  their  difTcreiitiatioii,  hs  would 
he  eNpeeted,  hciuii,  however,  antedated  l>y  that  of  the  cells  thai  form  the 
liiusciilar  tissues.  In  the  ^i>iiii<ii n,  lor  exain|ilc,  iiiilscle  cells  liecoiiii' 
(Icvcloiieil  fiom  .•miclpoid  epithelium  and  from  a  layer  underneath  the 
external  epithelium.  These  iiniM'le  cells  contract  slowly  so  as  to  cause 
o)icniiii:  and  i-hisintr  of  the  small  moutlis,  or  osciila,  on  the  surface  of 
the  s|)on!,'e  ill  ics|)onsc  to  movements  in  the  sea  water.  They  are  in 
depcmlciit  of  jiiiy  nervous  structures. 

Til  I'crtain  Cu  h  nti  niti  ^  the  muscle  cells  respond  a  little  more  (|uii'l<l\ 
than  in  the  sponges,  and  this  L'reater  efficiency  is  found  to  he  dependent 
upon  the  appearance  of  a  localized,  very  primitive  nervous  system' 
This  nervous  system  consists  of  six-cially  modKied  epithelial  cells,  o?- 
ritijiliirs.  scmlinir  hranches  from  their  inner  ends,  which  cither  come  in  i-oii 
tact  with  the  iniiselc  cells,  or  i  l)i<  tors,  or  li<'i-omc  interlaced  so  as  to  form  a 
network.  In  the  rcirion  Ijctweeii  tlie  recr-jttwrs  antl  the  effectors  tlie  v.''- 
w(irl<  at  first  serves  merely  as  a  structure  wlicreli\  the  entire  museiila 
tiirc  of  the  animal  can  he  hrouu'ht  into  harmonious  action   from  a  siiiiri' 


TIU:   K.VOI.I  TloN    OF  TIU:   NIRVOrS   KNSTI.M 


im 


,...5„-  on  the  surfn.-...  ns.  for  ..xnu.,.).'.  i"  tlu-  .•use  of  ,l.o  sou  nncnonc 

No  •'  of  |.'i«  jnl  >.    In  tl.-  j.-llvtlHh.  whirl,  i. ntraM  to  ll,.-  h.m.  a...-iuo„.. 

is  a  fr....  .novi..,  annnal    w  tin,)  that  th- pt-rs  a.v v  hi,hl.v  sp-.-al- 

i„.  an.l.  th..r..fon..  ...u-1,  .uor-  .....Mtiv...  an.l  that  th.  .n.,.nU..s  wln-h     ,.■ 
roooivo  «r..  .r«.>s„,itt.-.l  to  a  m-.ro  .lodnit..  n-rvo  notwork.  capa  .  o  not    nlv 
of  convovinK  th.  ..xoitatory  p. ss  fn.n,  on.  part  of  th.-  annual  to  anntlur. 


mjgpU^  Sp.n,e 


Sea  anemone 


Simple  form  In 
earthworm 


e  Addition  of 

055oci<Mlon  neurons 
inearltiworm 


but  also  of  in.printin,'  ...  tho  inipnl.-  a  .•hara-t.-ristie  rl.ythmie  adivity 
whi.  h  hrinjrs  ahont  th-  .-ntra.tion  o'  !,.•  h.-ll  an.l  th-  sunnnnn..'  n>ov.w..t 
„f  the  animal.  Tho  network  now  assuni-s  tho  function  of  an  n,J}U^irr 
as  woll  as  a  f  ransmittor  of  inipi  Isos. 

tho  n..1ju<tor  i«  an  rxtronL'ty  simple  stru.-t.nr,  an.l  it  is  poss.bU 


So  far 
that  tho  cfToetor  anil  rocopt.r  or 


IMS  a'o  (liii'otly  conniM 


ted  hv  fibers 


rnnnmn 


through    it.      When    wo    o. 


II10    SI  !liiii  ti 


till    inrirtihnitm 


m 


i  ' 


»n 


If-:     ,P 

ill   |. 


IM 


h 


7K4 


Till     IIVTIIM     \lll\iilS   >>>TI  M 


iff 


III 


oe 
b 


(huoIi  u8  tin-  ciirtliworni.  > mvlMi,  lolislcr.  clc  .  i    iniu-li   mnrt*  tiffliiitn  s|h> 

i-jnlizaliim  o{  llio  ailjiiNtti  < uis,  tor  imw   this  iiitrinii.liiitc  ni'ivmiH  ti« 

HUP  ht'i'omcs  rcilliTtfil   into  Mi-i-alU'tl   i/«jHf//i<i.  a   pair   oxwling   for  <'Mfli 

vf^'iiii'iit  ami  tli>-  variini><  |iair>  liciiiir  fciiiiii'ctcd  li>   (li'litiili- 

rii-rvr  Ntnii'luri'>.  ciin^iilutiMi.'  lln'  u'liiiulioii  cliaiii.     It   i^  in 

this  (.M'oiip  i(t'  animals  that   wr  have.  I'nr  the  lii>t  tilii"-.  ih'l' 

iiiitc  cvidi'iio'  of  till"  cxisti'iii'i'  III'  till'  III  iiniii,  whii'li  iiia\   If 

I'liiisidiTi'il  as  the  I'li'iiii-iitary  unit  ol"  which  tin'  ihtvous  s.\> 

ti'iii  of  all  till'  liit;hfr  aiiiiiials  is  Imilt.     A  iiciiriiii  iiiav  hi' 

fithor  st'iisdi'v  or  iiiutiir,  aiul  in  hotli  casoH  it  consists  of 

a  ('I'll   with  a  nucleus,  mio  loiii;  process,  called  the  ii.rnii. 

and    several    short    hraiiehiiit;    pr sses,   i-allcd    the    ih  ii 

ilrili!<.  The  ji\oii  in  its  course  ma\  ({i\e  nlT  a  liraiich. 
or  more,  at  riclit  aiiK'*'^.  these  are  sometimes  called 
riillnti  nils,  and  at  its  end  it  ma\  hreak  up  into  \eiy  line 
hranches  called  a  siniiiiisis.  In  a  sensorv  neuron  the  im- 
pulse is  transmitted  from  the  end  of  ihe  axon  to  the 
nerve  ci-ll,  wheri'as  in  a  motor  neiiron  it  is  traiismitti'd 
ill  the  opposite  direction  from  the  cell  to  the  end  of  the 
axon  I  Kiy.  '.Mi:',  r, 

The  simplest  arraiiKcmenI  of  sensor\  and  motor  neii- 
lons  to  eonstitute  fh(>  nervous  svstem  is  seen  ill  the 
eartliworin.  in  which  it  t"oriiis  the  simplest  t>pe  of  nlhr 
iirr  (Fin.  "Jltl,  No.  :>i.  The  sensorN  neuron  has  its  cell 
liody  in  the  skin,  and  its  axon  pioceeds  to  one  of  the 
seamen''  Kan^lia,  in  wliicli  are  laifje  ner\e  cells  whose 
thick  axons  jiass  out  from  the  irannlion  as  motor  lihers 
to  the  muscles  of  the  hody  wall.  The  dendrites  of  the 
motor  neuron  and  the  liranchinK  of  the  terminatiiui  of 
the  sensory  neui'oii  cause  a  very  line  interlacement  of 
norve  fihers  in  the  •.'aniflia,  I'ormiiiir  a  network  known 
as  the  HI  uriiftilf.  The  sensory  impulse,  on  reaehinj;  the 
!;an<rlion,  is  transmitted  hy  the  synapsis  to  the  den 
drites,  ])ri>hal)ly  witliout  the  (iliers  .icliially  joininL'  1o- 
srethei';  that  is,  the  neive  impulses  pass  from  the  one 
to  the  otlier  set  of  liranclies  hy  '•oiitact  rather  than  liv 
transmission   tliroiiuh  continuous  tissue. 

Hy  sueli  an  arranut'iueiit  it  is  evident  that  the  nervous 
apjmratus  in  eacii  seKment  coulil  cause  a  contraction  of 
the  miiseles  of  jts  own  nei|f)iliorhood,  liiit  that  a  stimulus  applied  to  one  r<'- 
eeptor  would  be  incapable  of  callinj;  forth  a  contraction  of  the  muscles  of  a 
far  <1istanf  sepment.  mneb  less  a  coordinated  contraction  of  the  musculature 


i;idin      i.l       l.tlV-'U' 
111II1I..I  ilivtfl.- 

l.rali  ;  a.  >ui'r.i 
i-.il.lijK<  ■!  nan 
ulicii;  /•.  siilx-. 
l.li.iKial  K.iiiKli"ii; 
.-      i-s.-pliami^     <<r 


iA^ 


Till     I  VOLITION    OK    THl     VKHVOIH   HVMIM 


78r> 


..r  th.-  whole  uniiual  muI.  as  woul.l  1.'  .•.•.,.....•.!  for  U. u.t.on.     U,  ..•udcr 

,|.is  poHHiM..  it  i. .-".v  tlmt  s ,u-a...  ■■,  ..onn.u.ni.at.on  Ik u.c  e. 

,..|.li.|,nl    l.-tw. tl,..    .l..T.T...,t    .,rM..Mt.l    ...M.'l.n       Tin.    ,s    ...r.-t.-.J    l.v 

auociation  n,uron»,  ca«h  of  wlm-h.  as  the  .uun.-  in.,,!,..,  ••on.mt,  of  a  uerve 

,.,.|1  with  it^  .l.n.lrit.  s  I,...nt.-I  in '-•Mi.di -ni  -I  an  .vn,         „-h  |..»- 

to  the  next  or  even  t..  nom-  ..."re  -listant  «M..Klion,  where  .t  end-  by 
.ynap.it.  The  important  ,.oi,.t  to  „ote  is  that  these  assoe.ut.on  ncurot.s 
do  not  leave  the  eent.al  nervous  systen.;  they  n.enlv nnoct   varn-us 

So  far  '       -nrRlia  of  eneh  segment  a.e  of  e-imil  .mp-tnoee.  but  . 
we  exr.       .    n    J.er  we  shall  fin.l  that   at  the  hea.i  end  of  the  nnimal 
,eve.    '  ol     ,r  .m^^'■ ,  *    -m..  fnse.l  fo^ether  to  for.,,  a  larger  «nn«h..n. 


^i;t 


I  ii(.  -''1... 


•--,:::/^"'tr".r"i  :vn"r!-^'  i;jr-n;'t:|-  'r^;};TV.:.:r 


.1,   jirttniK   filn-r;   i, 


which  lie.  just  hehind  the  .M.ll.t.  a>„l  f-o.n  ^^hieh  fihers  p.oceed  around 
the  gullet  to  unite  in  ficnt  "f  it  in  anothei-  laife  Ka"Blioi>.  which  usually 
shows  three  lobes.  These  laruer  L'anKlia  re-rive  aff-Ment  nerve  fibers 
from  the  eloselv  adjaeent  ini.nitive  sense  or-a.is  for  siv'l.t.  sound  and 
smell,  from  struetures,  that  is.  that  nre  really  highly  speeial.zed  reeep- 
tors  The  cells  of  the  retina  a.Ml  ...>•  hav  h.-n  made  .-aimble  of  reacting 
.o  impulses  of  lijjht  or  sound  !■  -ead  ..f  those  of  pain,  t..ueh  or  ten.p.'.a- 
ture,  to  whieh  the  reeeptors  ol  •  -e  int-KUment  ar.-  .speeially  seus,t./,Ml 
Thev  are  distnme  receptors  (pn.jieient  receptors.,  and  it  .s  evident  that 
the  nerve  reflexes  with  which  they  are  ..oncerned  are  of  a  higher  order 
than  those  loeated  in  the  se^Mi.ental  ^aiiKlia  themselves. 

Some  of  the  neurons  of  the  head  f;anf,'lia  are  tnerely  nu.tor  and  a-t  on 
the  muscles  of  the  head  end  of  the  animal,  hut  others  are  purely  assoca- 


-ir  I 


'I 


'i  If 


HI 


7S(i 


Tin;  (  TNiuM.  MKVdi  s  svsti;m 


'I  if 


\'U  I 


liiiii    iii'iii'oiis   ,'iikI   |)i I'll   iliiwii    tlir    t.'iiiii:liiih    i-li;iiii    to   tci'iniiiatr   liy 

syiiiipsrs  ill  iiiK'  in-  olliri'  iit'  llic  si';;iiii'iital  iraiisrlia.  Tlicsc  association 
iiiMifKiis  cxiTi'lsc  a  iliuiiiiiatiiiL:  inlliii'iu'c  over  liic  ai-tivitics  of  tlic  spfj- 
nioiital  i;an«lia,  so  tliaf  liny  iiia.\  lirfi'iiiiiiic  llii-  rcsixiiisc  of  llic  animal 
wlicn  its  sat'i'ty  is  tlui'.'iti'iii'il  liy  sdiih'  apjii   iirliint;  ciiciiiy.     Wlion,  for 

oxaiiiplo,  llic  stiiiiiihis  iiinilu !  iiy  sniiic  siu'iit  or  souml  of  an  ajiin'oacli- 

ititr  ciit'iiiy  is  riiTi\  I'll  liy  tlic  lirail  u'aML'lia,  these  will  tiaiisiiiit  inipnlses 
ilowii  the  L'aiitrliiiii  rliaiii  whieli  so  inlliieiiee  the  various  nerve  cells  of 
this  chain  as  to  prmliii'i'  in  all  of  them  a  eiioiilinateil  action  fur  the  pur- 
pose of  I'l'inox  ill!!  the  animal  from  ilaiiirer.  Kveii  sIhmiIiI  some  local 
stiinulant  lie  aetiiiir  mi  "Ue  or  more  of  the  se>rments,  the  response  may  be 
inliiliiti'il  on  aeeoiiiit  of  stimuli  meanwhile  transmitteil  liy  way  of  asso- 
ciation neurons  from  the  larsie  hcail  jraiiiilia;  in  other  wonls,  the  jiart 

eontrolleil   liy    the  si'miienlal   iraiiirlia    li mes  suliserx  ieiit    to   the   whole 

lliriiiitrh  the  ilciniiiiatins;  emitrol   of  the   lieail   !.'ani.'lia. 

This  illustrates  the  lieuinniiiu's  of  the  liili iiralioii  of  tin  )ii  d'oiix  si/xfi  hi  : 
ami  as  we  pass  to  the  stmly  of  the  liiirlier  animals,  we  shall  sec  that  this 
inteijrratioii  licconies  more  ami  more  eoiiiplieat"il.  and  that,  as  it  iloes  so. 
the  nerve  eeiitei's  aeipiire  the  power  of  storinir  awa.v  the  impressions  tlie.v 
receive,  which  they  mav  al'terwanls  appl.\'  to  re^'ulatc  the  retlcx  response. 
Thus  nil  iimnt  anil  roHlimi  come  to  tiiiil  their  place  in  the  nervous  inte- 
irratioii  nf  the  aiiiiiial.  The  alTereiit  stimulus  anivinsr,  let  >ls  suppose, 
at  nerve  cells  eoiitiollinir  the  movement  of  the  Icir,  ma.v  fail  to  cause 
a  response  of  the  e(iirespoiiiliii«r  muscles  liecausc  of  imi>ulses  meanwhile 
ti'ansmitteil  li\  assncialiiiii  neurons  from  hiirher  memory  centers,  for 
the  ;iiiiiiial  may  liaxe  Icnneil  liy  e\perieiici'  that  such  a  movement  as  the 
local  stimulus  \\iii,hl  in  itself  cill  forth  is  opposcil  to  its  own  licsf  in- 
ferests.  This  eN|ierii'Mci'  will  ha\c  lieen  stoieil  ;iw;iy  in  memory  nerve 
centers,  so  that,  whenever  the  locil  stimulus  is  ri'pcateil.  impulses  are 
iliseli.'ii  L'eil  fruni  the  memory  centers  to  the  local  nerve  centers,  iinil 
the  redex  respniise  lines  not  occur,  o?'  is  much  moililiiil  in  nature,  For 
storiuL'  awjiy  these  memories  anil  dtr  relateil  i>sycholotric  processes  of 
viilition.  ell'.,  the  aiilerior  )>iMtions  of  the  nervous  system  in  hiirher  ani- 
mals Ipccomic  very  hi'_'!ily  ilcvclopeil  so  as  to  constitute  tlir  hritin.  anil 
the  si'iiplc  chain  of  iranu'lia  of  the  iiivertelirates  is  I'cplaeeil  hy  Ihr 
spinal  rnnl. 

As  we  asceml  the  scale  of  the  \  ertelir.'itcs,  the  liraiii  liecomes  more 
ami  more  ilivelo|ieil,  until  in  the  hiv'hcr  m.'immalia,  siicli  as  man,  very 
few  reflex  actions  can  necur  inile]>enilently  of  the  hiirher  centers  Avliieli 
are  loeatcil  in  it  The  rellex  arc  now  involves,  not  one  nerve  center, 
hut   scMiiil,  anil   of  these   the  iimst    important   .-ire   loeatcil   in   the  lirain. 

There  is  thus  no  essenlial  ililTerincc  in  the  irencral  nature  of  inteijra- 


.i      ! 


liy 


for 


Uf- 


tiH 


it*'- 


use 


till 


nro 


1111- 
iiml 


/;.. 


Tin;  I 


Vl.l.l  THIN    OF   Till.    Nl.UVorS   SYSTIM 


r87 


lion  ill  the  nervous  syslciu  o 


f  llic  lower  as  eoinpjuc 


«1   with  llu>  lii«lioi' 


nnini 


lis,  lint  tliere  is  a  very  < 


listiiiet  iiioriilioloy:ic  .jilTereliee:  i 


II  the  lower 


,r  iiiverlelirate  aiiim 


Ills  the  i,'aii'_'li<'ii  nerve  < 


lain  IS  vei 


trill  to  the  aliinen- 


nnal,  whereas  in 


tarv  ( 


lakes  the  i)lae( 


the  hittlier  or  verl< 


lirate,  the  •^liiiial  eor<l,  whieh 


.f  th 


jranirlia,  is  < 


lorsal  to  the  aliini  ntaiy 


In  liotli 


:roin>s  the  hea<l  iiantrlia  ai 


lorsal  to  the  alini.ntary  eanal.  hnt   in  the 


vortehnites  these  heeoiiie 


iinieh  more  delinitt 


^tinetnie,  and  i-imstitiite 


the  l> 


This  inori)holo'j;ie  ililT<'i-.'net 


liftwceil  vertehrates  all 


1  inverle- 


hrntes  is  prnhalily  not  so 


riind; 


iiiienial  as  a 


t  first  siu'ht   it   may  a|>|><'iii-  <" 


be    for,  as  HasUell  has  shown,  it   is  possi 


Me  that  the  alimentaiy 


il  of 


the   invertehrates   is    real 


hoiii 


ilomiiis 


s)niial  eiiii 

inir  to  this  oliscrv 


1  anil  the  vei 
Ir 


it  rides  of  the  1> 
ihat    h 


IS   really 


ith    the    central    eanal    of   the 

1- 
f 


of  the  verteliiales.      Aeeor< 
hapiiene.l   in   the   latter   -rr""!'  " 


limals  is  thai  the  u-ani-'lia  have  t'lowii  ii|) 


as  to  siirronn 


il  the  alimeii- 


tar.v   eanal   anil   so  eo 
eanal  heiinr  meanwhile  piov 
f  ventral  <lo\\  nu'iowth  of  t 


nistitnte  a   i-on 


tiniions  striietnre,  a   new 


o 


ilinieiitary 
i.le.1  hy  tin'  enelosnre  .if  a  spaee  as  a  result 
1,..  l,o,iv  walls.     Althon-rh  this  view  has  not 


lieen  general 


■  pted  liv  hiolo-risls.  then 


no  inherent  reason  v 


hv  it 


should  not  be  aeeepted.     It  is  no  more 


to    11' 


dered  at  than  the  w 


■11- 


Unow  n 

]>assay:e  from  ai| 


•t    that    a    new    ri'spiratoiy    system 


b( nil's    develo|M'd    in    thi 


natie 


to  land  ainphihiiin''* 


The  (iliers  i 


,f  the  sensory  neiiroiis  in  \  ei 


tebrates  are 


[•led   tiiliether 


to  form  the  jtush  mif  nm 


//,<  of  the  spinal  eord.  and  the  e< 


I'll   bodi 


,f  th 


neurons  a 


re  located   not   on 


th 


>nrface,  as  in   inver 


tebrates.  but    in 


th 


osterior  root  "janji 


li.l.  the  cells  beinc  ineelei 


1  to  the  libers  by  T-sha])e<l 


junctions. 


The  olfaetory  nerve  is  the  only   one  i 


11   the  hiulicr  v.'rtebrates 


ihieh  retains  its  primitive  e 


(indition. 


In  tl 


le  vei 


Icbrate  iinimals  the  s).inal  memli.i 


ill  the  inteirralii'ii 


■  f  llii 


central  nervous  system  is 


the  motor  nenron 


the  fibers  beili!.'  collected  in 


the  aiihriiif  rix'ls.   Toward  the 
not  only  from  the  seirment  in 
soeiatioii  neurons  from 
central  nervous  s\  slem. 


if  this  nenron  impulses  are  traiismi 
ihich  it   is  its..|f  located,  bnt  by  way  of 
f.ii    distant   parts  of 


tted. 


ither  scLiiients  or   Iroiii 


In  otl 


icr  Willi 


Is.  th 


iv  motor  neuron  ma 


th- 


V  t  ransinit 


Isos  which  cause   t 


impii 

nieiits.  which  are  coon 


he   iiiiiscles  to   peilorm 


linated  with  th"' 


.f  ad 


local    independent    iiiovo- 
iid  which 


jaeeii 


lit  scLrnients  a 


may 
appropri 


be  of  widclv  varyiii-.'  ty|>es. 


The  motor  neuron 


has  therefore  very 


itel 


V  heen  en 


lied  the  fntnl  I'lniiiioii 


fuilh.  and  it  wi 


11  b. 


mam  i 


b.jects  later  to  show   tl 


le  conditioi 


IS  umler  w 


if  our 


ihich  several   difTerent 


comiietiny;  intluenees  m 


iiv  obtain  |losses^ 


ion  I 


if  this  Jiath 


I    «• 


ter. 
aiii. 


Ta- 


1' 


1 1' 


I  II  \ni:i{  I, WW  II 


THK  rUol'KKTlKS  ol'  KA(  II  I'AIM   <•!'  TIIK  UKIhKX  AKC 

Having;  l)riclly  traced  tlif  |)li\  sinlngic  (li\  cldiuiii  lit  of  tlio  ncivous  sys- 
tem, we  are  piTi)ai<(l  to  ('(iiisidei-  in  fjieater  detail  tlie  luculinr  function 
"f  paeh  of  the  parts  wliieli  enter  into  tiie   foiniation   "f  tlie   i-eflex  nrp. 


Hi  ^:i  !| 


'.  I' 


THE  RECEPTOR 

With  the  advance  in  animal  oitiaiii/ation  is  associated  llie  development 
of  tlic  ability  to  api>rcciale  and  discriminate  !■  'ween  external  phe- 
nomena, special  orf^ans  called  leceptors  lieiiiir  e\  'vcd  to  reecive  the 
stimuli  which  these  occasion.  Those  receptors  \s  ..eh  are  distrihuted 
over  the  skin  of  the  aninud  are  called  (.rlirnnl  or  i  rtt  run  jilors.  and  ari' 
especially  adapted  to  react  to  such  stimuli  as  temperature,  ])iessiire. 
and  pain,  hut  at  the  tore  end  of  the  aniinal  eci  t^m  irccptur^  liccoinc  in'O-c' 
hiphly  specialized  so  as  to  react  to  stimuli  coniinir  from  a  distance 
that  is,  to  stimuli  tliat  are  not  produced  hy  contact  of  external  olijects 
with  the  suriaec  of  the  nninnil.  These  specialized  receptors-  -sometimes 
called  prDJirii  lit  include  the  eye,  the  ear,  and  the  olfactory  epithelium. 
Receptors  are  also  provided  in  the  iutfM'ior  of  the  oriianisni  ("r  the  pur- 
j)ose  of  reeeiviiiir  stinuili  dependent  ujion  the  activities  of  the  organism 
itself.  They  may  he  ealh'd  niUri'nl  /■  ri  jihirs.  and  we  nniy  further  dis- 
tinguish two  trronps  of  them  namely,  those  which  come  fintn  the  sur 
faces  (if  till'  mucous  memhianes  and  those  whicli  <'oiiie  iiom  the  suh 
stance  of  the  various  oi-jiaiis  aii'l  tissues  ihemseKes,  as,  for  example, 
from   the  sulistance  of  muscle  <m-   tiiidon. 

A  receiitor  may  !"•  ilefineil  in  a  'jeneral  \\a\  as  a  nnchanism  in  whici; 
some  particular  kind  of  stiiindus  pruducev  chanL'es  that  result  in  the 
excitation  of  the  nerve  lilwr  with  wliich  the  receptor  is  cioiriecti'd,  al- 
tliouirh  the  vtnuulus  in  itself  is  ineapahle  ,,]'  e\eitin!.'  the  nerve  tilier  lu 
other  words,  as  Siieriiniiton  ]»uts  it,  the  receptor  has  the  threshnid  oi 
its  exeitahility  raised  to  every  kitnl  of  stimulus  save  one.  toward  wliich 
it  is  lowered.  A  nerve  fiber,  for  iiistani'c,  lesjuinds  to  every  kind  ot 
stimulus  appriixiuiately  e(|iuilly;  a  receptor  will  also  I'cspond  to  these 
same  stimuli,  Init  with  s.'reat  ine(|iiality,  since  each  recept<u'  is  specialized 
to  react  to  one  kind  of  stimulus  and  to  others  only  when  these  are  very 
strong. 

788 


Tilt;    PHOII.HTIIS    OF    I:A(  H    l-AKT    OF    THK,    HKKl.LX    AK« 


789 


I,   i.  oftrn  a  .Uni-ult  ...altrr  t..  .lotormin-  just  exactly  what  it  is  in 
the  nature  of  thr  stimulus  that  makes  it  onpahlc  of  affoctin«  one  reoepto. 
niHl  not  another:  for  example,  it  is  often  merely  a  question  ofthe  ra  e 
,.f  vil.rnti..,,   of  the  stimulus.     I.iirht    nn.l   heat    rnys   are   both   due   to 
vibration   nf  the  ether   whieh    tills  spae..     When   these   vibrations   are 
sl,.w    thev  slirnulat..  reeeptors  tha.   have  been  speeialize.l  for  appreeia- 
,ion  of  t.'.n.peratnre.  but  vvh-n  they  are  rapi.l  an.1  ex'st  as  rays  of  l.prht. 
,1h.v  no  lon.'.r  niTect  tb..  teM.pe.nture  reeeptors  but  only  the  hichly  spe- 
,.inii/e.1  ree..,>t.,rs  .,f  the  r.tinn.     Similar  vibrations  of  the  air  in  plaee 
„f  the   ..ther   eause   snu,„l    r.n.l    stimulate    th-   auditory    reeeptors.     Tt    is 
,,„ite  likelv  that  tl„.  v..,.,.pt,.vs  in  ,lilTerent  irronps  of  animals  are  aftune.l 
,.,  veaet  to  .liffrrent   rates  of  vibration.     For  example,  a   oat  ean  hear 
hipher  pitehe,!  notes  than  man,  ami   it  is  possible  that  tne  retmas  of 
M.me  animals  r.s,,„n,l  t,.  rays  vibratin-.  vvitb  a  .lifTe.vnt   fre,,neney   fron, 
,l,os..  to  vvhi.h  tb,.  n.tina  of  m.in  is  a.lapte.l     In  this  ronn-etion  ,t  ,s  „f  m 
lorest  to  note  that  the  tonch  reeeptors  of  the  shin  respond  so  promptly 
to  stimulation  that  one  hundred  vibrations  of  a  tuning  fork  per  seeond 
,..,„  ,,..  f..l,   as  s<.,.ar..t,.  stimuli,  ^^^u■u■.^  to  the  ear  at  thw  freqneney  the 
fork  emits  a  eontinuons  note.    The  reeeptors  of  toueb  are  therefore  more 
prompt  in  their  response  than  the  reeeptors  of  the  auditory  nerve. 

\Vl,en  onee  the  reeeptor  has  been  stimulated,  the  impulse  passes  and 
is  transmitted   to  the  nerv.  eenters.  where  it   is  translated   '"'" J* J"'- 

ti,.„lar  sensation       Tl ondition.   are   r.:my   not    unlike   those   whieh   ob- 

1,in  in  the  ease  of  the  various  pbysieal  instruments  used  to  receive  and 

.'.Hvert    into   tl I.-tri n-rent    MinH.li   nf  heat,   li^rht.   -henneal   enersry 

,.,e  The  reeeiver  .e-iuired  to  brinu  about  this  transformation  must  be 
espeeiallv  ..onstruef.d  in  ea,-h  ease,  that  for  lieht  beinc  the  aetinometer. 
,|,at  for'nn.ti.m  ,1,.  dvnan.o.  that  f..,-  h.a.  the  tb.n.H.pile^  ami  that  tor 
ehemical  ener-.v  the  eoneentration  eell.  Each  of  these  phvsieal  instnv 
ments  nmv  he  "eonsidered  as  a  speeiali/ed  receptor  for  the  purpose  of 
„n.duein..'  an  eleefie  enrrent  out   of  other  forms  of  energy       _         _ 

Tn  aceeptiim  the  ;d.ove  analogy  «e  must  not  fail  t„  bear  in  mmd 
„,„  verv  feeble  siinudi  are  often  able  to  set  in  operation  nerve  impulses 
tl,,,  ,n.'  as  potent  as  those  pro.lueed  by  nnwh  stron.-er  stimuli.  Here 
ncain  -ve  have  a  phvsieal  analogue  in  the  ease  of  relay  currents,  m 
,vhieh  a  feeble  eleetrie  eurrent  mnv  operate  to  complete  the  eireuit  from 
independent  sourees  of  electric  diseharee  and  thus  set  in  m-tion  a  mucti 
larger  amount  of  enertry. 

These  .'cr.eral  eonsiderations  of  the  nature  of  a  receptor  naturally 
lead  us  to  the  law  of  thr  .prrifir  prnprrlir,  nf  vcrvf.  which  is  to  the 
effect  that  however  ex.-ited.  eaeh  nerve  of  special  sense  gn-es  rise  to 
its  own  peculiar  sensation.     Thus,  in  whatever  way  the  ehorda  tympan, 


^^MH^&^ 


"•(» 


Tin;    CINTHAI.    NlltVDI  S    SYSTllM 


1"      1 


ncrvp  is  stinnil;iti'il  ( rluniicnll.v ,  incrlinnii-iilly  m-  elect  lic.ill.v'i  <Iiii-iMi;  its 
l)nssiiL'e  iici'iss  tlie  tviii|i:iniliii,  tlie  si'iis.ilion  evnked  is  that  "f  t.'isie. 
And  M>  with  tlie  leeeptur:  wliiitever  the  ine;ili>  li\  wliieh  it  is  excited, 
Avhethei-  by  tlie  ii.-irticiil.ir  kind  of  stiiM\i!iis  I'lir  which  it  is  adapted  or  It.v 
e\cessi\e  iiileii^it ies  <il'  other  stiniilli.  e\cit:iti..ii  always  evnkcs  the  same 
sensatii)ii  It'  the  uptic  iieive  or  retina  is  nieehanicaliy  stimulated,  as 
hy  |iiessuri'  a'.'ainst  the  miter  eantlnis  of  the  vyv  or  hy  an  electric  enr- 
reiii.  the  sensation  is  that  of  liirht.  Applyini;  these  facts  to  less  well- 
known  receptors,  sin-h  as  lliose  of  heat  anil  cnld,  it  is  interestinjr  to  note 
that  stimulation  of  a  "cohi  spot"  hy  extreme  heat  or  liy  inechMnical 
or  electrical  stiiniili  hrinjrs  out  the  sensation  of  cold. 

Properties  of  Epicritic  and  Protopathic  Receptors 

A  valiiaMc  iiniiii>iii<i  nf  i-<  n  iilnr^  nf  !!■!  nkiii  has  lieen  deiiionsti-atcd  hy 
Head  and  his  ]iui>ils  li.\  esperiiiients  on  himself.  Head  found  after  sec- 
ticiii  of  the  skin  iier\es  of  the  radial  nerve,  for  example  that  deep 
])ressure  and  ]>aiii  were  still  present  in  the  area  sii|>plied  hy  the  nerve, 
indicatiiii.'  that  these  deep  sensations  are  carried  1>.\  the  sensory  (ihers 
])resent  in  the  iiiiiscnlar  nerves.  In  >in'h  a  paral.\/ed  sensory  leuion  tin' 
])o\\('r  of  general  locali/alion  is  fairl.v  trood,  altlioiitrh  li'-dit,  touch,  tein- 
]>erature  and  sii|)erticial   i>ain  are  I'litirely  ahseiit    in   the  ovcrlyinir  skin 

In  the  CISC  of  the  lin'.'ers  the  nerves  of  deep  seiisihility  iiin  in  the  ten- 
dons of  the  liiiLTer  ninscles,  so  that  aftei-  sc\ci-;ince  of  thi'  ciitaiieons 
nerves  and  temlons  of  the  hand,  all  seiisihility  is  l'oiic. 

Hiiriii'.'  the  resicni'iation  of  the  cut  nerve  the  cutaneous  seiis.-ilioiis  re- 
aitpear  :  ;  two  periods:  one  uroup.  called  the  iinihiinilliii-.  hciriiis  to  ap- 
pear ill  from  seven  to  twenty-six  weeks,  whereas  the  other,  called  ( jmrilii-. 
does  not  full;-  appi'ar  for  one  or  two  vear>.-'  The  protophatic  sensations 
arc  of  a  distim-lly  lowt'i-  order  than  the  epicritic.  When  ihev  alone  are 
lU'cseiit,  there  is  the  sensation  of  jiain,  hut  not  that  of  fine  touch;  tem- 
]>eratlire  seiisjitioiis  ;ire  t'elt  when  extreme  deirrees  of  heat  or  cold  ahovc 
:!S  ('  or  \\<-\\<\\  -in  ('  are  applied  to  the  skin,  hut  not  for  slight  de- 
jri-eis  ;  the  powi  r  of  ilisiiiiiiinatiii'/  hetwecn  two  points  is  almost  cntirel\ 
ahsent  ;  and  the  sense  of  locali/.at ion  is  \ery  iniperfect.  i''or  example,  tlw 
person  uill  ..fieii  refer  the  \i'H\\\  tli.it  li.is  actiiall>'  liien  stimulatccl  to  ;i 
iici'_'hliorin'_'  iioniial  portion  oi'  skin.  l*rotopathi<'  seiisiliilit.v  is  more  or 
h'ss  distrihiiled  in  spots.  .-Hid  it  is  stroimly  "alTective"'  in  character,  caiis- 
ill-r  an  illlelise  sllh.iecl  i\  c  sriisal  loll       .\  si  ill  I  111  lis  that  causes  onl\    moderate 

pain  under  noiinal  c lilioiis  pioilncvs  in   ;■   '•protopathic  arci"  a   |iain 

that  ma.v  lie  intense 

The  epirritii-  sensation.  .is  will  le  inlerreil   from  the  fori"/oiie_'.  responds 


Tin;  i'U< 


,n.inii;s  i>y  i;a<ii  i-\kt  ok  tiii.  iii;fi.i.x  Mir 


7!tl 


,..  ,l„n.  ura.l..s  ..f  sti.nulnf ion.     lU  it  vv.  .-an  iVcl  the  l.ul.t.'st  t..u.-h_a.  .1 
ran  .lisciininalc  tl..  li...  ^1  u'.a.Us  of  f.np.'.atu.T  l-tw.'...  2..    an.l  ...     ' 

Tin-  ,...w..r  ..!•  LM-ali/a,.. f  liu-  sti.nulus  and  tlu-  alni.ty  t..  .lis.-vn.Mnat.. 

lK'lNn-.-n  tMo  points  also  iHuru  uith  ..pl.-ritic  n-.-n.Mation 

"l:   ,„..  spinal   .-.nl  tlu-   n.-v.   .i.KTs  .-a.-ryin.   on.   /,/..     "f  -nsafon 

„,.,.  .M-on,..-.l  to.„..-1  !....•.  in  tlu.  s.ns.-  Ilu.l   pain  s.-nsations.  wi-tlM-r  .l-'op  o, 

;.ro,;;path„-,  mn  in  ,1,-  sa„..-  -ohnnn  in  tlu nl.     Lik.-w.s..  ....np.Tatun. 

sensations.  nvIiWIi.t  |,rotopatlii.'  or  .•pi<'''i'i'--  ••""  t-J-'-th.-r. 

The  Peculiarities  of  Bach  of  the  Separate  Sensations 
Temperature.     Tl,..  n..-..,.tors  for  ,..n.p.-tnn.  an.  armnu^Ml  in  ..-nn.ps, 
so,no  ImTo.  s..ns„i.-l  U.  heat,  others  fo,-  .-.,1      These  ^-'-^^  "^ ^^^''^'^'^ 
,„.^.  ,,„.,.l  ,„,„  „„,/  rnU  .,,.,t..     Thev  ean   he  verv  eas.lv   .leteete.l  on  an 


! 


v:  1 


f 


sV 


l-ij:     _'nl,       riMMi..i,-lli. -i.t^if 


,.,va  o,  skin  hv  n.-ans  of  a  pointe.l  hollow  vessel.  lh,ou..h  uln.-h  water  is 

„u.,le  to  tlow  at  a  I- ratu.v  a  little  h.low  or  ,   liul-  ahove  that   o        e- 

skin  The  instrnn.-nt  is  ..alle.l  a  ther,no..sthesion....er.  On  a  part  ot  the 
.kin  where  there  are  no  heat  an.)  -ohl  spots,  the  .l.-rnio  ..s,h,.Mon,e,er  will 
Hi..it  MO  sensation  either  of  heat  or  „f  .ohl.     This  i,  .har...!  '■'-'"  '•';';- 

,,,,win,  of  the  par.   as  a  neutral  -,-<.     A.   otl.-r  pi s  „    will  eal  l.o.tli 

a  sensation  of  heat,  iii.ii-atin.  the  presence  of  heat  spots  or  a  others  a 
sensation  of  eohl.  in.liiat.n,^  the  piesen.-e  of  enl.l  .po,s  1.  «  i  I  ..•  >.o.,.,l 
that  certain  of  the  spots  are  mn-h  n.ore  reae.ive  than  others  anil  that  tlios, 

of  eohl  are  n.n-h  th-  r.  nuiii-i  o,.    ^e  Im..  -iu:,..     I'.o.i,  heat  a.el  .-n 

spots  are  most  fmp.ent  a.  the  nipples,  th-n.  in  onh^r.  -on.e  the  ehest    th 
nose,  tlie  anterior  surfa-e  of  the  arm.  an.l   the  ah.hnuen.      I  hey   are  leas. 


nose, 
ni 


arke.l  on  tin-  expos...!  snrfaee  of  ,iie  skin,  su.-h  as  .he  fa..,.,  ami  tli.y  aiv 


1    ■• 


'•I  *     r 


If-:  111 


pu! 


702 


TIIF   ••FNTRAI.   NF.BVOI'S   8TRTEM 


■  'iii 

i 

'   n  . 

:■•■        |!:^       ' 

, 

li;-- 

^1 

% 


also  very  infrequent  in  the  Rcalp.  They  are  almost  absent  from  tho  mucous 
membranes,  whieh  explains  why  one  is  able  to  swallow  a  liquid  that  is  too 
hot  for  the  hand. 

The  nriitnus.^  of  llic  temperature  sensation,  as  with  all  the  other  cu- 
taneous sensations,  depends  very  mueh  on  the  condition  of  the  skin, 
Itoinp  most  sensitive  when  this  is  at  the  ordinary  temperature,  but  very 
imperfect  when  it  is  either  very  Imt  or  very  cold.  There  is  also  very 
marked  tidaplnlion  of  the  sense.  This  can  be  very  well  shown  by  tlie  simple 
('Xi>crinicnt  of  taking  three  vessels  of  water,  one  at  a  moderate  tempern- 
tun',  one  very  hot  and  one  very  cold.  If  a  finder  of  one  hand  is  placed 
in  the  hot  water  and  a  fintier  of  the  other  in  the  cold,  and  they  are  left 
there  for  a  shi»t  time,  until  the  skin  has  assumed  the  same  temperature 
;is  the  water,  and  then  transferred  to  the  lukewarm  water,  the  fingfi 


.1 

I'iK.  ;n'.— CiM  '■I'nH  (A)  anil  lu-.1t  »lnls  (/I  I  '.f  an  area  nf  iskin  nf  the  right  h.nml.  In  .  a.  ii 
r.ise  the  ni'.it  intt  ?i^c  vfn«.ttinn«  wtre  f  xi-ericiuTii  in  the  hlafk  areas,  less  intense  in  the  hne-i. 
.in<l  least  in  the  il'itli  il-  The  Mjnk  .irt-.ts  rti  rf^f  nt  p.irts  where  no  special  sen>alion  of  eith<  r 
kind    'A.is    (  xjieri«  n(  eil.       (I'rnir    (ii.l.In  h<-i'lir  * 


!'    .U' 


transferred  from  the  cold  water  w  ill  feel  hot,  and  that  transferred  from 
the  hot   water  will  feel  cold.     Temperature  sensation   also  produces  a 
iii.ii  ki'il   iifi\ifirt    iilli  f-i  fli  I  I .      Thus,  if  a  cold  coin  is  placed  mi   the  fori' 
head  and  tlien  removed,  the  cold  sensation  will  persist  for  some  time  in 
the  areii  of  skin  on  A\hieh  the  coin  w as  laid. 

That  the  receptors  for  heat  and  cold  I'cspond  only  to  one  kind  of 
stimulus,  or  if  to  others,  only  when  these  are  excessive,  can  be  well  ii 
lust  rated  by  the  experiment  of  touchintr  a  coW  spot  with  a  very  hot  ob 
.iect:  the  sensation  will  be  that  of  cold.  The  hot  ob.jeet  has  so  pronounced 
;i  power  of  stimulation  that  it  has  overstepped  the  threshold  for  heat 
of  the  coid-adapted  receiptors.  The  sensation  of  cold  is  elici'ed  moi' 
promptly  than  that  nf  \v. •ninth     The  distinction  between  a  warm  and  .i 


THK   I'KOl'KBTIES  OF    KAIH    I'ABT  OK   THE   RK.KI.K.X    ARC  79^ 

hot  bnth  may  rc.llv  depend  on  the  fact  that  in  the  Iftttor  the  ooM  spots 
nrp  stimulntr<l  lis  well  M  those  ..f  heat.  It  is  at  Ici.st  intorostinj?  to  note 
that  the  phvsi..l..v'ic  reflexes  stimulated  by  either  a  cold  or  a  very  hot 
Irnth  are  the  snino:  thus,  a  rise  of  l.lood  pressure  and  h  .Mmtraetion  of  the 
tnuseles  of  the  skin  oeeur  in  both  eases. 

The  Touch  Benie.-  In  order  to  investisate  the  t-u.-h  sense  neeurately. 
v„n  Frev  has  d.vise<l  a  method  of  using  hairs  of  different  thiekncss  ea.-l. 
.nounted  on  a  different  handle.  The  hair  whieh  prndnoes  a  sensation 
of  f.uK'h  when  pressed  on  the  skin  so  that  it  just  bends  is  then  similarly 
pressid  on  one  scale  pan  of  a  Imlanee,  and  the  weiirht  required  in  the 
..fl.or  s,.i,le  pun  tn  hold  the  beiun  horizontal  when  the  hair  just  bends,  is 
nseertained.  From  the  dinmctrr  of  the  hair  onr  emi  then  ealeulate  how 
manv  jmnns  per  s.piniv  n.illinuter  are  neeessiuy  t..  elieit  the  sensation 
(.f  loueh.  The  following  quantitative  results  l.,-.ve  been  obtained  by  ap 
plyinir  von  Frey's  method  to  different  pints  of  the  body: 

Om.  per  (if|.  iiim 
T<iii;;iio  and  nosio   .  .      

I'lp    

FiiiKcr  tip  linrl  forohpii.l ^ 

Hack  nf  finccr '^ 

Palm   " 

n  s 

Knionrm 

n-.K-V    of    hniwl '■-' 

Ciilf,  shnuMor '" 

.■Mxlnrnpn    " 

Outdido  nf  thiuli -'' 

Shin  and  snlo   - 

Tlaek  .'f   fr.rrrirn- ...  

T.oin    *" 

Th;it  til.'  sons.,  of  toueb  is  locMtod  in  spots-  tourh  ■>potK  can  best  be 
,bMnonstrMt..,1  ..„  tlie  calf  of  tl,..  lep.  If  this  is  shnvod  and  then  earefully 
explored  with  a  fairly  stiff  hair,  it  will  bo  found  that  there  ■•ht  onlv 
some  twelve  to  fifteen  spots  in  nn  arra  of  a  square  centimeter  at  wbieh 
•lie  b.iir  •■Ml,  l.c  f.lt.  Metwcn,  these  spots  there  is  no  snisatb.n  ot  tou.-h. 
That  these  spots  are  composed  of  specialized  receptors  can  be  very  clearly 
shown  by  i.ivssinp  :i  fine  needle  into  .me  of  them,  when  no  pain  will  be 
experienced  but  only  n  i>eenlinr  shotfy  sense  of  pressure 

(^areful  exnminati..n  -f  the  p..sition  of  the  touch  spots  wi)l  further 
show  that  thev  inv  iiroup.'.l  around  hair  follicles,  particularly  on  the  side 
from  which  tiie  hair  extends  the  windward  side,  we  may  call  it.  This 
f;ict  explains  the  >ve11-kn.."n  experience  that  an  object  may  be  felt  more 
acutelv  on  a  iiairv  surfa.-e  lluui  iift.r  that  surface  has  been  shaved.  Th.' 
hairs  i...nd  sliphth    when  the  obj..!   romes  in  cmtaet   with  them,  thus 


!   ; 


ft-     1 


li 


I  '  il 


if' 


7;iJ  Tin;  i  intiiai.  miivoi  s  systim 

caUHiliir  iji-csmiic  to  lir  exerted  i>ii   llie  luiir   fiillieles,  so  tllllt    tlio   tnueli 

(•(irpnseles  in  llie  iieit'tilioili 1  <'t'  tile  lollieles,  or  i)erli!i|).s  tlie  fliip  nerve 

jdexus  whieli  siirroun.Is  llieni.  l.ei e^  .xeite.l.     The  influence  of  liiiirs 

in  inereiisinn  tlie  loneli  sehs.ition  eaii  lie  'ieinonstnitetl  l>y  tlio  voii  Fiey 
niellMMl:  for  ex.iniple,  in  one  ex|>eiiinetil  o\ei'  iin  area  of  !1  si|UMre  mil 
liiiielei>  of  skill  Willi  liiiiis  |. resent,  1  niilliuranis  w.-re  found  to  pro.lii<-e  tlie 
sensation,  ulieleas  alter  llie  liaiis  lia-l  lieeii  reinoveil,  it  r<'i|uiretl  :5ti  liiilli 
cm  Ills, 

Tlie  fie(|neiiey  of  toueli  eorpnseles  ili(Tels  very  liiueli  in  .lin'i'relit  Jiarts 
of  tlio  l.ody.  They  are  most  plentiful  on  the  (inireis,  relatively  infre<|iioiit 
over  the  skin  of  the  haek.  ami  very  searee  in  tlir  skin,  direetly  over  lioii\ 
surfaces.  They  are  entirely  ahseiit  from  the  eornea,  the  eoiijiinetiva 
of  the  upixr  lid,  and  the  -jlans  (leiiis.     The  adeipiate  stimulns  for  loueh 

is  evidently  deforniali f  the  siirfaee.      Pressure  exerted  over  all  the 

toueh  eorimseles  of  a  ))ortion  of  skin  is  iii.l    felt      This  can  1>e  demon 
st rated  liy  dippiii'-'  the  liniri'r  into  niereiiiy.     The  iiressiire  of  the  iiiereui\ 
is  felt   on   the  siiifaei'  hut   Hot   in   the  siilMiier'.'ed  ))ortion  of  the  finirer 
Toiieh  is  the  most  responsive  of  nil  the  sensMtioiis     Thus,  as  has  alreadv 
heeii   noted,  a   tuiiinir  fork   ean  he    felt    vihr.-itiii'.'  h.\    the  liliu'cr  when   t" 
till'  ear  its  note  is  a  eoiitiinioiis  one.  ami  the  sliniiili  produced  liy  a   v 
volvinir  serrated  wheel  ean  he  felt   hy  the  tiii'.'ers  as  sepal  ate  even  u]! 
to  a  rate  of  live  or  six  hundred  stimuli  per  second.     Adaptation   is  also 
a  marked   feature  of  the  touch  sense,  as  is  the  experience  of  every  "in- 
mIio  has  wDin  llannel  niiden-lolhiiii;  or  a  plale  of  false  teeth. 

Closi'iv  related  to  the  tactile  sense  i>  the  jiowcr  of  ilisirhiihiiitlon  hi 
liitni  Inn  i>iilnls.  This  is  tested  liy  liiidiii!,'  at  what  distaueo  the  two 
lioints  of  :\  )).iir  of  calipers  sljind  in  order  to  he  distintiuished  as  separate 
The  'esiilt  in  any  uiven  pail  of  tli.'  Imdy  varies  a  little  aeeordinir  to 
whether  the  iioinis  rest  on  touch  corpuscles  and  accordiiiK  to  the  rela 
tioiishi|t  of  the  calipers  to  the  li.iir  folliih's.  <>n  an  averajio,  however, 
we  may  take  the  follow  inir  distances  in  millimeters  as  heinir  those  at 
which   the  two  points  ean   he  distintruished   over  different   areas  of  the 

hody: 

mm. 

Tl|>   111'    toiii;!!!' 1.1 

V..l;ir  ^nrf;  •■i-  ,,(  (in-,  i    li|. 2.r. 

Il.,rs:il   ,.f    tii>l    |,I,:,1:mi\ C.'i 

l';.lin   ..r  lian.l II.". 

link    ..f   li.'Oi.l :"•!.'! 

V.-A.'\i  of  M-.-k <5ld 

Mhl'llc  of  liiic'li.  ii|i|'('r  .11  III  iiii'l  -I'll' fi'.l 

Tt   is  clejir  from   this  list   that   the  power  of  discrimination   tends  to 
iliniinish  in  proportion  to  the  lessenin;;  mohilit.\  of  the  part.    It  is  greatest 


<  I 


Tin:  rm.innns  ok  i m  ii  rxirr  ..k  tim    iiin.i  \  ah. 


7!tr» 


„  „,..  tip  of  tl...  I,mun..  .m.l  .1...  ti,.  of  fho  Ma-o-s;   ,.   .s  l.ns.  on    l,c 
,,,,,„iv..lv  innnol.il..  .kin  of  .ho  l.nok.    Tlnse  .l.stan.vK  nr.  .mu.;1.  Uss.h 
„.,  ,„.|Mls  n.^.  on  ..„  lo,M),  .orpus,.!,..    m.lrr  ,h.  s-  <-on.l.t.ons   for    n- 
„„„....    tho  .liMMn...-  for  tl...  volnr  si-l,-  of  tl,..  fin^.T  t,p  or  ..von     or     , 
pnl,n  of  tin.  Inn,.!   .n.      !-    .'^.ly  on.-t..ntl,  ..f  a  n.illMn.t...- ;  nn.l   for  tin 
„nn  Mn.l  l.n.k  it   ,■  .ouo    -nln.-.a  lo  lu.lf  n  .nillin...tor. 

/.,„.„;;:„(.■.,.  ./  is  ..  V.MV  ......nrat.   pro.-.ss.  at    Las.    ,n   tl,o  n.ost 

s,.nsitix..  parts  „f  tl in,  l-nt  n,.v..Hl,..!..ss  it  i.  v.ry  prol.nbly  «  ma^- 

t,.r  ..f  ...l.l-a.ion      An  ..u.l.u....  .,f  this  is  tin-  fa.-l  tl.a.  -n  tl,..  n,.,.!,  .„...■ 
Ulllv  sp,  riali/..!  ...tina  tl,..  p-nvr  of  ln..ali..ation  of  ol.J.-ts  n,  tl.o  v.sual 

fl..l,l  is  a  p,.......ss  .,f  .■.hn.a.i .O'l  -M..- i.'.,.-...     For  this  ,.as.,n  a  K.-son 

f,.nn.  .1 •o.,.....i.al  .a.an,...   has  |.....n   .-.• -l.  -,  not   1...'..  o  thr 

ohjoots  ul,i..h  In.  -..s  until  aft..,-  1,..  has  l.ar.,...!  hy  his  .xp.  n.n....  of  t..  . 

♦nst..    .t....  t..  a.s,„ia...  .h..  po.-.i f  tUr  r-tina  s.,n,-.lat...1  w,.h  a  ...r.a  n 

,„,,„r  Un.  xis,n,l  li.M      ir  .hi-  is  .n,..  for  .h,    i.-.ina,  i.  »  alsn  proha  1.  y 

r,,..  lor  Ion,  I,     Vu-  U .s  .sp-rin,.  n.  of  Aris.o.l,.  is  ..xp  n.a  1.1..  on    h- 

sa.H.  hasis.     If  .1...  tinK-s  a,.,  .-r.-ss...!  an.l  a  n,a,l.l..  p'  .....llMt^  .■.■»    h. 
..n.ss...l  fi„..'.-s.  i.   ^^ill  In-  .VI.  as  .l..uhl.-.  sinr,.  n.m    it  ton,.h..s  two  skn. 

s„rra....s  «hi..h  hav..  n...  lr,u  , nst.na...!  to  t..u..h  the  sa„n.  ol,,,..-!,    -nt 

,.l„..ato.l  t..  f...l  .litT.n.nt  ..h.i.-ts.     INp.ri..,,...'  asso-iatrs  th.-s.  two  sk.n 
aroas  with  .lin-..n.nt  .,l..i.'..ts.  no.  with  .h..  same  ..hj.'.-t. 

The  Pain  Sense.     I.  uas  a.  on.,  ti,,,..  thon-.'ht  that  th..  s,.n>a.,. I  i^an, 

was  .In.,  t..  .,v..,sti„,nlati..n  ..f  any  kin.l  ..f ptor.  hnt  it  is  nnw  known  that 

for  this,  as  f..r  ..thor  skin  s..nsations.  th..ro  oxist  spo.-ial  m.opt..rs.    Thus, 
it  is  f..nn.l  tha.  in  .-..rtain  pa,.s  of  th..  ho.ly,  MU-h  as  th,.  .....n.-a,  an.l  t..  a 

,.,.,tain  ..xt..nt  in  th..  -lans  p..nis,  pain  r,..-..pt.,rs  al-n..  ar..  p....s..nt,  an.l 

in  ,lis.-as..  th.^  s..nso  ..f  pain  .„ay  h..  ..ntirely  al-.-lish.-.l.  chorea,  that  o 

,.„„.,,  ,„,„,;„..  ,1,1.  ..„n.li,i..n  h.-in.  .-alL-l  anal^.sia.     Ov..,s.,nu,la.>on  ., 
a  t.,n..h  spot  .l.u-s  n..t.  as  wo  havo  soon,  oanso  pain  hut  ..nly  n  sense  of 
pressure.     Alth..u,^h  pain  is  appn'oia....!  hy  spooial  r...-..p.ors.  .he  .-harao- 

lor  of  tho  pain  is  ,Iop..n.lont  .m  tho  ...hor  sonso  r. pto,s  s,„.wl.an,...nsly 

..x,.i,o<l-  r..r  .xan.i.lo,  a  thrnhhinc  pa.n  is  .luo  t..  tho  si,„ultan..ons  pres- 
sure pro.lno...l  l.v  .lila....l  hh-.l  v.ssois.  oto.  A  s,.„sa.ion  of  pan.  a.T.Mn- 
pnnios  .-ortain  r.lhxos  ..f  a  p,.....olivo  natu.o  .  n..oio..pt.v..  ,-...lox..s,  pau-.. 
s-..-.,  a,„l  «h,.n  ,1,..  -vllox  is  ahson,  th..  pa,-,  is  likoly  t..  suOVr  .lania.-...  On 
this  aeeonnt  the  pain  nerves  nn.y  he  re._'ar.l..l  as  tr,.pl,i..  nerves      The 

s..nso  ..f  pain  n.ay  als nr  in  struotnr.-s  ^^U\,■h  a,o  .lov,.,.l  .■!  ..nl.nary 

sonsil.ility,  Mi.-h  as  tho  iutostino  an.l  .In-  unt.r 


ih 


Ui 


■I 

:  3 


r-1 


i! 


riiAi'i  :i;  i  xxwiii 

Tin:  I'Uol'KIMII  >;  or  l   \<  ll  I'.MM  nlTIIK  WKI'f-KX 
Ali'i     •  •  Hill  M 


V 


THE  NERVE  NETWORK 

III  mII  MiiilM.il*  iil"n  1-  lilt   1  'I'li'i.l.iiit    ~.  no  ilii.'il  |ir..|..|.lii>iiii iiiiiinii ' 

exists  liot\M'rn  tho  \;iiii>iK  lu'iironx.  il  .-  trjm^inissiun  i.f  flip  iu'Vvr  iiiip'iKt 
ilfpcinTmir  '111  I'niifiirnity  infhrr  )li;iii  e..    timiitv   cf  llic  ijiini     ts  tliiit  cdii 
sfitiitc  tlic  roflcx  arc     Tliis  tfnii^iii»sii)ii  iii;i>   !>.■  itTci'tfil  tlnonph  n  <«,vtt 
,ipsis  poiniiie  in  fKHtm-t  riflior  \'     li  dnndritf^  i<v  w  ifli  iicrv .'  coIIm      It  i- 
fxtrrnu'ly  diffifiilt  to  l;n<nv  \\\u  tlitr  '''fic  is  really  nny  nnntomif  con 
fitniity  liotwmi  tlu'  \nri(iiis  filtnrs  \(  '    'h    t'"iiii  tlir  iiclxMir'k  in  tho  irrM 
mattrr  nf  tho  i-ciitrnl  iutvoik  -;y«;ti  ni      W"  shnl!  nut  ,ittniii>f  tn  dispii  — 
tliis  \c\o(l  qiii'stinii  liiiT,  liiit  in  (iiiliT  tli;it  \\c  (iia\   li'Mi'ii  «iiTtiotliin(j  nf  tl  I 
possihlo  fiitiitioiis  of  a  nrrvo  notworU,  \\o  may  con-i'l' r  that  proscnt  in 
'lie  \\  alls  oi'  till'  iiiti'stiiic  'pltAiis  ol'  AiiiM-l.at-li  ami  M<-      ner."*     Tliis  plr\' 
spptiis  to  liavp  an  important  fnnotion  lo  pcrfonn  in  i  onncction  witli  tit.' 

■iiMntiTJc  r.'llc'x  -Ml'  oaL'c  4t''li'.      At  'last  it  |i::s  1 n  sliown  lt,\   Afocir  tli.i' 

aftor  tran'J'scrtifin  nf  tlio  intostino  tlio  tniis.-iilar  and  rpitholinl  stniofnros  1)0 
comt'  n'L'cMorntt'l  considoralily  cailior  tlian  llir  nTvous  ploxns.  bnt  tliai 
tlip  niycnforic  rcdiN.  wliidi,  it  \\ill  lie  rcinonilx-rcil,  is  cliaraetpri/ril  li\   .i 
wave  nf  inliiliitimi  preer.liinr  ono  of  eotiti'aetinn  <1no«  not  oepiir  nntil  after 
tlie  p1e\us  liiis  lireii  rrtrnirr.vtrd 

NETWORK  ON  SKIN  NERVES 

A  very  imi)ortaiit  t \  po  of  iiervo  network,  from  tho  medical  vic\vp<iint. 
is  that  which  is  produce"!  eloso  to  tlioir  receptor  endinsrs  hy  tho  branch 
ins  "f  til*"  afTeient  fliiors  of  the  skin.     Tliron>rh  these  branches  the  vas- 
iMilnr  reactions    rolbnvlntr  the  applieatinn   of  an   irritant   to  the  scnsoi'\ 
s\ii(';iee   t:ike  )iiaee   u  jtlioiit    tlie  ilit  er\  elil  ion    of  any   nerve  cells.      Tt   Us.ij 
'.I  I.e  llioiiulit   that   such   reflex   vasodilatation  de|)eii«led   n|ioii   the  trai  ~ 
mission   nf  an   impulse   alontr  an   atTerent    neuron   tn  an   eflfcrcnt    vas" 
dilator  neuron,  a  view  stiii  tly  in  eonsoiianee  \>  itli  the  ne\iron  hy])otliesi- 
That  such  is  not   tho  cast  .  however,   is  shown  hy  the  fact  ohsorved  h.v 
\inian   Piiire«   that    irritants   sueh    as   niustnrti    oil    applied    tn   tho   sUii 
or  cornea  etintinue  to  produei'  their  iisiial   reaction    for  somo  time  aft' 


i      I  I 


i  !!-  •   i 


TH( 


llti.riRTlKH   OK    K\iH    lAHT  !>►     IHI     HIH.I  X     Mt' 


I'M 


«,ction  of  the  iKmliTior  rouln  of  tl.c  s,,i,.ul  -onl,  l.ul  fu.!  t..  do  ho  ,t 
,l„.  wvsv  lil..T>  «1-..  .lit  ..n.l  «llo«.a  to  ,lrK.-.u,af.  .M  ,1'  tin  sl.nn.h  a.r 
l.looked  bv  npplvine  .-."•nino  to  tt.e  skin.  Wl.at  actually  happon.  .^ 
..vul..ntlv  llmt  tiu-  i.npulM  s.t  up  by  li,.-  inituut  a.  it  truvols  ....  i1m- 
ulT..,..-nt"(n..M-  pans...  ....  to  ....c  -f  tl...  Mmioh.  s  «»...n.-  .•'■t.r.v.l  to,  «|o..« 

„M.-1.  it  tl..-.  pr 'Is  to  tb..  I.I00.I  v..,.,.lH.  ulMd.  it  ■•....-«  to  a.li.t.v 

Tl.at  su.-h  vav,.lilato,.  i.np.i'MS  ...ay  l-r  t.'a..sn.ilf.l  -lo„„  th.-  Iib.'.'s  ol 
.,u  a//rr.»(  ...•rv  l1a^  Im.m.  .■o„lin....l  !«>  Hayliss.  ulu.  fo...,.!  that  vaso. 
'.Hlatntion  o..,M,.ml  i.>  tl.o  l.i.ul  li.'.l.  ^^  l.-^n  tb."  P-Ht.-.m.^  sp.nal  .o..t> 
u.p'  >.tiiiiiiliit.'.l      >«••■  p'lU'-  -•!♦ 

Post  root 


thr   »kMi   i«   tr.in'milli  .1    I'V   tlv    •■  ■  - 

1  ar'    ■■!    it    1  .i--ii'i;   '■•■    1'.'     !■  ".!»■  ■  il 


ri        A    •' 

iiir 

1  ,*  aiMl"-!   '" 

.IIH  t.. 

Ir.li    il 

!(n-    spin.l 
..11- •    !'■ 

.III 

In    tins   pniph.Tal   iMan.-lii,.-'   .>f   ^h'■   ^iHV.n.t    lii-.r-,   of   tlu'   sUm,.    >^ .; 
have  tlHMetorc  a  so.t  of  >uu.op,!o  ^l.i.h.  liUe  ll.at  of  ec.tain  for.>.>  oi 

,-..i,.,„„,...,.s  ,s..,.  ,..„•  7>^ .  is  ,^„-l■^■  "'■  -'-i"^  ''^  - 1-:';^^^'^  ';"■  '.'"■ 

'ra„s„.ission  of  a  so„so,-y  imp..lso  to  a,.  olT.-<-tor  oriran   w.tl.m.t   the  .n 
..rvonli....  of  .,-.-vo  ....lis.     S,...l.  a  .•...l..v   is  k„..w..  as  a.,  ax.m  n.tl..x.  a...l 


it  is  ..vido..t  that  it  ...ay  ........r  tl...o...-'li  .^ny  lil-T  whi.'l.  j-mvos  otT  bra...!.,  s. 

,,„,.  t.-av..li..«  to  a  M.„s....y  su.'f. Tl,..  ..tl,--  t,.  s.,,,,..  ..tT....tor  orp...,   a^ 


ill  ih 


.UMstric   liciv.s  to  til''  l.l.i.i.l.T      ■"■<■   |.,-i!. 


THE  SYNAPSIS 

At  the  iM.i.it  of  ......ta.-t  b.tu......  a  i.ia....h  of  on. u.-o..  and  a  nvvx. 

,.oll  of  the  n.-xt.  «..  bav..  s....,>  ,liat   tli-r..  .n-'s  m  >>.-•'"-.■  ><""^;;'  - 
the  synapsis.     Altho.iv'b  this  is  ,l..s.Tib...l  by  bistoL.g.sts  as  a  tuft-l.k.- 


,i    t 


MICROCOPY    RESOIUTION    TEST   CHART 

ANSI  and  ISO  TESI  CHART  No    2; 


1.0 


I.I 


12.5 


;-iiiiiM  _ 


c  a 


2.0 


11-25  IIIIII.4 


1.8 


1.6 


^     APPLIED  \M/\t3E     Inc 


''6;  *8;      QiOC      P.'o.-.e 


m    r 


:uni 


i  W  ■: 


m^  ^^ 


!IS 


Tin;    (T.NTKAI.    NKKVOIS    SVSTI.M 


liijiiicliiii!:  iif  tlic  I'lid  (if  till'  iixoii  (Fiu'.  -tlTi,  it  iiiiiy  rciilly  consist  of  a 
sort  of  iiiomliniiio  -the  synaptic  iiiciiiln'aiic.  It  iioniiits  the  nerve  im- 
pulse to  11J1SS  ill  one  diicctioii  only,  from  syiiai)sis  to  cell.  Of  what  this 
mciiilii  may   he  coiniiosed,   we  do   not    Unow,   'nit    there   can    he   no 

doulit  .  to  its  frreat  functional  iinportance  in  connection  with  the  in- 
tciriatioii  of  the  central  nervous  system;  for  examiile,  failure  of  an  im- 
pulse to  pass  lietweeii  two  neurons  may  he  due  to  retraction  of  the 
synaptic  iiiemhrane  from  the  cell,  or  to  alteration  in  its  iiermealiility  to- 
wards the  nerve  impulse,  iicrhajis  ;is  a  ciinsei|uence  of  ehaiiKcs  in  surface 


\'lg.  Jll,".  Aril"!  i/ 111""  "f  n.llati  im1-  Ir.Mii  tin-  |..-.Uriu 
pMslrriiT  li.rii  -).  ;i'»'niiii«  tilitr  in  (...vUrii'i  .nluiiit.^;  H. 
li ,    syiiap^i^.       ll'i.iin     Kanmii    y    C'aj.il.  I 


r.c.l    liliir^    annul. I    the-    iiH-    "f    tl" 
i.llalrral>;    C.   y.  Il>   of   |..i»uri.ir   li.nn, 


li  1  fi^f 


tension.  Similar  chanjres  may  also  he  hrouj.'lit  ahout  l>y  tlie  action  ot 
electrolytes  or  liy  chloroform,  strychnine,  and  other  drills.  As  we  shall 
see  when  we  come  to  study  tlie  rellexes  of  the  hi'^'her  animals,  there  can 
he  little  doiiht  that  it  is  in  the  synaptic  meiiihrane  that  many  of  the 
peculiarities  i-eside  which  charaeteri/e  I'oiiduction  in  a  reflex  arc  as 
compared  with  that  in  a  nerve  trunk.  The  phenomeiiii  of  summation, 
of  reciiirocal  iiihihition,  of  facilitation,  etc.,  are  uiidouhtedly  depend- 
ent upon  such  alterations.  The  synapsis  is  also  almost  certainly  the 
seat  of  fatiiriie  in  the  central  nervous  system,  and  it  is  possilily  the 
structure  whose  piiysiolo<;ic  activity  heeomes  upset   in   surjrieal   shock. 


, 

! 

Hi' 

i 
j 

III 

l^ 

Tin;  rK(M'i.i{Tii;s  ok 


i:.\<ll    I'AWT   OF   Tlir.   Kr.FI.LX    ARC 


79!) 


THS  NERVE  CELL 

Aside  from  l.ci.i!;  a  inoctini;  pliu-o  ..f  fil.ors  .-...nin-  from  various 
sources  the  nerve  eell  miiy  have  other  fui.etioiis,  sueh  as  that  of  reiu- 
fcreiiis;  impulses,  .just  as  a  relay  ...ay  reiufoi-ee  a-,  eleetric  eur.-ent.  It 
is  also  .•esi.onsii.le  for  ii.aiMtaiiuMjr  the  .iutriti(m  of  the  axon  with  whieh 
it  is  eom.eete.l.     It.  the  ease  of  the  posterior  root  til.ers  of  hi-her  am- 


Fig. 


Normal    nil    fr( 


:inl<Ti..r    linrii.    staim.l    to    >h"W     Ni>-l's    Branuk-s. 
iImo:ii     llowrll.l 


mals,  this  fuiietio.i  is  pi'olmhly  the  most  impo.-taiit  which  the  cell  per- 
forms, for  it  has  l.eei.  fo.i.id  liy  separating  the  frauiilin  fr(.m  their  blood 
supplv  in  the  fros:  that,  althousrh  the  cells  de-enei-.te  in  about  two 
veeks,  sensory  impulses  continue  to  be  t.-ansmitted  thi'ouj:h  the  Ran- 
plia.     Similar  observations  have  been  ii.ade  in  the  case  of  the  cab,  ui 


which  the  eell  bodies  o 


f  the  neui'oiis  lie  on  the  surface  oi 


the  ganglion 


!        ! 

1    . 


;'■  1 


11 


J' 


I  - . 

\ 

l\ 

,    1 

'.  »'. 

■■' 

i  ' 

■■ 

1 ; 

I   !  1 

I'll  ■ 


I    ! 


1'  It 


i.l,! 


11  .I'M 


'If " 


I'l^'. ' 


'    il'l 


\     \ . 


!        '    I 


f^     I 


!3  ;  - 


8(H) 


THF.   CKNTKAI.   M'RVOUS   SYSTllM 


mass,  from  v  Lioh  they  ciiii  hi'  separated,  Icaviiig  merely  the  iieuropile, 
through  wliieh,  however,  tlie  reflex  contiimes  to  be  conveyed.  After  a 
lime,  of  course  in  t)iis  case  also  the  I'etlex  disappears,  because 
an  axon  can  not  live  indclinitc].\-  alter  It  lias  iicen  separated  from  its 
nerve  cell. 

These  facts  regardiiif?  the  j-eneral  function  of  the  nerve  cell  arouse 
our  curiosity  as  to  its  morphologic  sti'ucture.  When  nerve  cells  are 
fixed  and  stained   in  various  ways  they  show  certain  elements  in  the 


^ll     ,:■« 


-3  H  /^f-j  . 


1  ll^    -""       I'.-iit  of  ail  a:iti'ri(ir  Cfirmial  eel!   frn;n  thr  c;ilf's  <iiinal  c<inl,   staiiuil   tn   sliiiw   TUurntitirils. 
ov,  axon;   a.   h.  r,  .icmlritis.      ( I-'roni   r.ttlic.l 

cytoplasm— namely,  (1)  large  granules  or  masses,  which  stain  deeply 
with  basic  dyes  and  are  i-alled  N'issl  bodies  (Fifj.  '2i)S\  and  I'J^  a  line 
tietwork  of  fibrils  passing  through  the  cell  substance  from  one  process  oi- 
(b  iidrite  to  another — neurofibrils  (Fig.  20!)).  These  ai)pearanees  in  fixnl 
and  stained  preparations  are  possibly,  however,  entirely  artificial;  for  when 
nerve  cells  are  preserved  in  a  living  state  -by  being  suspended  in  some  of 
the  aiumal's  own  lymph  or  bluod  serum  it  is  found,  when  they  are  ex 
iiiiiined  by  the  aid  of  the  Mlti'MiiiiiMiiscoi)!'  i  see  pjiiic  .'I'i ) .  thiil  the  eyto|ibism 


Tin: 


I'lton.iniis  (IP  r.Acn  i-aut  of  tiii;  ri;ki.i:x  auc 


801 


is  ('(mipoM'.l  uf  ii  visrnns  llui.l  full  of  cKtnMiM'ly  niiimte  };iiiiiul.-s,  <"ii.-h  of 
which  jipiKiivntly  consists  of  a  colloi.lnl  solution  summiKh'd  l.y  a  lipoid 
envelope  (Fisr.  -M(t).  When  the  tenip'-'fitnre  is  raised,  the  j;paniiles  dis- 
appear, and  wlien  the  cells  an-  deprived  of  oxy-en,  the  cytoplasm  ami 
,n,el,.us  !....• swollen.  A  similar  swellinj;  of  the  cell  and  nucleus  super- 
venes upon  sec'tioM  of  the  axon:  and  in  stained  specimens  the  Nissl 
granules  disappear  and  the  ,,rotoplasin  stains  dilTusely  (chroniatolysis). 
"  In  onil.rvoi.ic  life  the  pro.-csses  of  the  nerve  cells  appear  to  be  capn- 
bio  of  un(ier-:oin<:  a  certain  amount  of  aiuehoid  movement,  and  fd)ors 
jinnv  .mt  from  then,  indicating  therefore,  that  in  the  development 
of  the  iiervons  svstem  the  nerve  cells  appear  tirst,  and  the  nerves  suh- 
se.pientlv  -row  out  from  them  to  their  proper  destination.  Prolifera- 
tion of  isolated  tissue  cells  in  rilro  has  been  observed  for  many  other 


y 


tlif    ].r"tiililaMi!.      I  I'r.im    M.ii  iiumci.  I 

tissues,  such  as  cardiac  muscle,  renal  epithelium  and  connective  tis- 
sue. Its  occurrence  indicates  that  the  therapeutic  i.rineiple  that  the 
aim  of  treatment  should  be  to  -ive  the  diseased  or-an  a  rest  so  that  by 
cell  regeneration  it  may  rcover  its  lost  function,  is  one  which  may  ap- 
ply to  the  nerve  tissues  of  youny;  aninmls.  Whether  adult  nerve  eells 
may  reirenerate   is  as  yet   not  certain. 

This  s;ro,vins;  out  of  nerve  fibers  from  their  .-ells  is  the  essential  na- 
ture of  the  development  of  the  nervo\is  system  in  the  dcvelopin-^'  animal. 
At  birth,  unlike  the  cells  of  other  tissues,  those  of  the  central  nervous 
system  are  already  provi.led.  N..  new  ones  are  add.-d  during  postnatal 
life.  The  axons  irradually  deveb.])  from  this  inherited  stock  of  nerve 
eells,  and  bv  eonneetinsr  with  oth.'r  neurons  .servo  to  brlnj:  about  the 
intcM-ation  which  is  the  imi><'rtant  chara.-teristic  of  the  adult  nervous 


ft- 


\l 


802 


THE   CliNTRAI.   NI:KV()1  s    SYSTIM 


systfiu.     The  iiioit  complex  tlic  intcffrntioii.  the  higher  tlic  iiitelligeiuM' 
of  the  animal. 

Besides  i)erformiiiK'  these  fuuetioiis  the  nerve  cells  serve  as  store- 
houses for  memory  impressions,  certain  types  of  them  beliiy;  espeeially 
adapted  for  this  function.  The  diiTcrcneos  observed  in  the  relative  thick- 
ness of  the  cell  layers  comp<>sii:sj  the  cerebral  cortex  arc  more  or  less 
associated  with  the  function  wliich  it  can  be  shown  the  dilTerent  areas 
of  this  possess.  N<  ve  cells  are  extraordinarily  sensitive  to  deticieiicy 
in  oxy>;cii  suiiply,  iiiiil  yet  little  evidence  of  oxv «( n  couNumptioh  by 
the  brain  can  be  revcalcil  by  the  nsnal  methods  of  investigation  (  pajie 


niHiK 


THE  INTERMEDIATE  OR  INTERNUNCIAL  NEURON 


It  would  be  jirotitless  at  this  stai;e  to  cmisider  the  possible  inlluelices 
that  the  intermediate  neuron  may  have  on  the  impulses  i>assins.'  alonfr 
the  reflex  arc.  Before  doing  so  -wo  must  see  how  the  problem  can  be 
approached,  for  it  is  i)laiii  that  the  nenron  in  the  case  of  the  simpler 
reflexes  is  too  short  to  make  any  investifration  of  its  jteenliar  functions 
a  possibility.  We  must  study  the  characteristics  of  some  type  of  re- 
flex in  which  this  neuron  is  drawn  out,  such  as  the  scratch  i-cflex,  in 
which,  as  we  shall  see,  it  extends  from  the  shoulder  area  of  the  cord 
to  the  lumbar  region. 


*',  '■ 


iri 


^i   1 


CHAPTKlt  I, XXXIX 


TlIK  RKFLKXKS  OF  THK  Sl'INAIi  ANIMAL  AND 


SPINAIi  SHOCK 


TTavins  bcpdmo  familiar  witli  the  pofuliar  proportios  of  each  of  tlio 
sti\iPturrs  wliipli  ffo  to  make  up  tlio  roflox  arc,  wp  may  now  proceed  to 
coiisidor  the  function  of  tlie  arc  as  a  wliolc.  It  may  l)e  well  first  of  all 
to  consider  briefly  the  experimental  method  by  which  such  studies  may 
be  made.  The  object  aimed  at  is  to  simplify  the  conditions  as 
much  as  possible,  for  it  will  be  evident  that,  in  the  intact  nervous  sys- 
tem, with  the  brain  excrcisinj;  a  dominatinir  influence  over  the  great 
ma.iority  of  all  the  reflexes,  it  would  be  impossible  by  ai)i)lyinff  a  given 
stimtdus,  to  predict  exactly  what  kind  of  reflex  response  it  might  call 
forth.  The  rencx  will  be  conditioned  upon  the  accompanying  influence 
whi(di  the  brain  exercises  on  the  reflex  involved. 

Tn  order  to  render  the  reflex  unconditioned,  we  must  remove  the  in- 
fluence of  higher  centers.  This  ciin  be  done  experimentally  for  the  re- 
flexes of  a  great  part  of  the  body  by  cutting  the  siiinal  cord  above  tlic 
level  of  the  segment  in  which  the  reflex  inidcr  investigation  resides. 
f=!ome  of  the  reflexes  elieitablo  from  the  cord  isolated  in  this  way  in- 
volve only  one  or  two  lu'ighboring  segments,  whereas  i.'liers  spread 
over  several.  The  reflexes  which  have  been  most  extensively  employed 
are  those  which  involve  the  musculature  of  the  hind  limbs.  Since  some 
of  the  recci)tors  concerned  come  from  the  skin  of  the  flank  and  shoul- 
der areas,  the  section  is  usually  made  at  the  upper  end  of  the  thoracic 
region  of  the  spinal  cord. 

Spinal  Shock  in  Laboratory  Animals 

Immediately  after  the  operation  a  profound  condition  of  depression  sets 
in,  involving  all  the  reflex  arcs  in  the  separated  portion  of  cord.  This 
condition  is  known  as  .«/>(>k//  shork.  It  suitervenes  in  all  classes  of  ani- 
mals having  a  spinal  cord,  but  is  much  more  profound  in  the  higher 
than  in  the  lower  animals.  As  a  result  of  this  depression,  the  part  of 
the  body  below  the  section  exists  in  a  limp  and  flaccid  condition,  and  the 
ai)plieation  of  even  very  strong  stiiriuli  to  the  skin  will  evoke  no  form 
of  reflex  movement.    In  the  case  of  the  lower  animals,  such  as  the  frog, 

son 


n 


i! 


m 


sn4 


Till'.  <i:\TK.\i.  \i;kv(11  s  systkm 


I  Ut 


I     I' 


J.     1^ 


nji  ,  u 


f 


f 


■•i     i 


tlio  ciniditid.!  l)OKiiis  to  pass  ofl"  in  I'lmn  twcniy  niimitPH  to  linlf  nn  liour, 
after  wliicli  a  stiiiniliis  applifil  to  tlic  skin  of  tlic  foot  is  followed  Ity  n 
tvpici'l  llexioii  inovenieiit  at  knee  and  liip,  tlie  so-ealled  (lexion  reflex. 
In  tlie  rat)1)it  very  little  reflex  response  is  elieitaMe  for  several  hours 
after  the  operation,  Imt  in  a  few  days  the  reflexes  return  eoniplotely 
lielow  the  level  of  the  seetion.  In  the  doj;,  on  whieh  a  srn'at  deal  of 
work  has  heeii  done,  the  involved  rejrions  of  the  liody  are  profoundly 
paraly/ed.  The  skin  is  in  a  more  or  less  unhealthy,  unnatural  eondi- 
tion,  the  surfaee  cold,  the  haii's  niPlled:  and  if  eare  is  not  taken,  the 
slijrhtpst  alirasion  of  the  surfaee  uuiy  result  in  a  nasty  ulerratinn.  On 
neeount  of  the  paralysis  of  the  eentevs  of  niieturition  and  defecation, 
there  is  also  incontinence  of  urine  and  of  feces. 

The  Reflexes  in  the  Spinal  Animal 

With  reasonalile  attention,  however,  the  dog  makes  a  wonderful  re- 
covery. After  an  interval  of  two  weeks  the  hind  limlis.  although  com- 
pletely paraly/ed  so  far  as  voluntary  movement  is  concerned,  hcgin  to 
show  eonsidenilile  si<rns  of  imi)rovement.  The  first  reflexes  to  return 
are  those  cone,  ed  with  the  deeper  structures,  such  as  the  vascular 
reflexes,  thus  lirin-rin};  the  skin  hack  to  its  normal  temperature  and 
condition.  The  reflexes  of  micturition  aiul  defecation  also  soon  return, 
so  that  the  animal  no  lonjicr  suffers  from  the  continuous  discharge  of 
urine  and  feces.  Aliout  the  same  time  the  kiirr-jirk  liecouics  elieitahle. 
This  reflex  is  obtained  hy  tappin<r  the  tendon  Mhieh  connects  the  patella 
with  the  tiltin,  the  response  lieinir  a  smart  contraction  of  the  extensor 
muscles  of  the  knee  .joint.  The  ffc.rwn  rrflr.r  also  hejrins  to  reappear. 
This  is  elicited  hy  applyins;  a  pinprick  or  other  hurtful  stimulus  to 
the  skin  of  a  lower  extremity,  and  when  fully  developed  consists  in  a 
flexi(.n  (.f  the  knee  and  hip  joints.  The  evident  oliject  of  this  move- 
ment is  that  the  stimulated  parts  may  be  removed  from  the  source  of 
stimulation,  and  it  is  plain  that  all  stimuli  that  produce  the  flexion 
7-eflex  are  such  as  would  cause  in  the  intact  aninud  a  sensation  of  pain. 
Such  stimuli  are  thus  classified  as  nocuous,  and  the  reflex  is  styled  a 
vodcriitirf  r(  flr.r.  Accompanyinpr  flexion  of  the  stimulated  limb  the  op- 
posite or  contralateral  lind)  usimlly  under^'oes  a  definite  extension, 
called  the  croxxnl  r.rliuxion  rrflcr.  The  occurrence  of  this  toftether  with 
the  flexion  of  the  stimulated  limb  is  an  important  thin<j  to  remember 
in  testinsr  the  reflexes  in  man.  Malinsierers  who  attempt  to  make  it  ap- 
pear that  they  have  some  lesion  of  the  spinal  cord  may  know  that  if 
such  lo^ion  exists  no  nioveiiictil  of  the  le<r  ocei-rs  when  the  skin  is 
stimulated,  but  they  are  unlikely  to  know  that  under  these  eonditi<ms 
the  opposite  leir  also  fails  to  show  a  simultaneous  extension. 


RKKI-K.XKS    IN    TlIK   SI'INAI.   ASlMAf.    AM>   SI'INAI.   SIKXK 


805 


That  the  ii.M-iccptivc  ivdcxvs  slmuld  U-  nuwuff  tlu-  first  to  ictuin  nftcr 

spinal    ti-a..s,.r1i..n    is  <.t    si.l.Tal.ir    i.itn-cst    as    ii.diratini:   thnr    itn- 

poitaiKM-  ill  tlu"  pn.tf.-tic.n  of  tlic  animal   from   injury.     Tlicy   aro  tlio 
css.'iitial  rellcxcs  of  (U'tViiso,  aii.l  it  is  consi.l.Tahly  later  in  the  rc-ovory 
of  tlie  animal  l.cforc  itH.-xos  <lrpcn.lcnt   upon  stimulation  of  oilier  tac- 
tile  reeeptors  Lejiin   to  show   themselves.     The  most    important   of  this 
latter   un.up   of   more    s|)eeial    rellex    movements    ineludo    the    so-ealled 
serateh  reflex  and  the  extensor  thrust.     The  svralrh   nlhr.  as  its  nam'- 
implies,  is  the  seralehinif  movement  of  llexh.n  and  extension  of  the  hind 
limb  at  a  rate  of  ahout   four  eontraetions  p.-r  se<-ond  that   oeeuvs  when 
a  meehanieal  stimulus  is  applied  to  the  flank  an.l  shoulder  area  of  the 
animal.     Tor  example,  if  we  «ently  draw  a  pencil  or  ti.e  finsfers  haek- 
M-nrd  and  forward  amonit  the  hairs  on  this  rettiou  of  the  spinal  annual, 
the  e..rresi)ondinir  hin.l  limb  will  be  brou-ht  up  so  that  the  elaws  are 
approximately  at  the  i)la.-e  stimulated,  and  the  limb  thus  direeted  will 
undertto  n  scries  of  flexions  and  extensions,  desijrned  evi<lently  for  the 
purpose  of  scratchinir  the  area  ..f  >kiii  that  has  been  stimulated.     If  the 
stimulus  is  a   weak  .me.  only   the   initial   stages  of  the  movement   may 
oecur.  such  ns  the  iireliminary  flexion  of  the  W<!.     As  we  have  already 
stated,  the  receptive  stimulus  calli- -   forth   this  reflex   is  very   spcciflc 
in  nature.    A  pini>rick  or  roii(.'h  friction  of  th.'  reflex  area  will  not  ,)roducc 
it.  nor  will  the  application   of  heat  or  of  a  siiifrlc  electric  shock.     The 
most  ade(|uate  stimulus  is  one  siiiiulatintt  as  nearly  as  possible  the  con- 
dition which   would  be  |)r..duccd   by  the   movement   on   the  flank  of  the 
animal  of  m.iiic  insect.     This  more  or  less  eomiilicatcd  scratch  refl.-x  can 
of  course  also  be  elicit«>d  in  animals  whose  spinal  c.u-d  has  not  been  cut. 
but  we  can  not  i>redict  in  such  cases  whctlu'r  the  rctlex  will  occur.     The 
brain  mav  inhibit  the  reflex  arc  and  prevent  the  ni<.veiiiei.t.    In  n  spinal 
animal,  however,  the  reflex  always  occurs  provi.lcd  an  adcpiate  stimulus 
is  applied.     The  irrcat  importance  of  the  serateh  reflex  in  the  study  of 
the  phvsiolosrv  of  the  spinal  cord  rests  in  the  fact  that  a  laiw  stretch 
of  cord  is  involved  iii  the  reflex  path.     The  afi-erent  imi)ulse^;  must  enter 
at  a  much  higher  level   than  the  cfTcrent   impulses  leave,  and  between 
these  two  points  there  must  exist  a  lout;  intraspinal  neuron  (see  page 
8i:5).     This  permits  us  to  study  many  conditions  inthiciwinj.'  reflex  action 
which  otherwise  in  a  reflex  located  in  one  se-iiieiit  only  it  would  be  im- 
po.ssible  to  investi<;atc.'' 

Thf  rrffiisor  thrust  is  elicited  by  ai>plyin<;  pressure  to  the  pad  of  the 

paw  or  the  sole  of  the  foot.     It  consists  of  a  .juick  extension  movement 

of  the  correspondiu};  limb  usually  with  a  flexion  of  the  opposite  limb. 

Ai'ter  coiii|>lclc  recovery  from  shock,  the  paralyzed  ].arts  of  th<-  body 

arc  capable  of  performing'  even  more  complex  movements  than  those  al- 


H(l(! 


Tilt;    CKNTRAI.    NI;HV0I!»   hVSTKM 


Ml*     I 


•  u 


i  ■'iliilil 


i    f! 


■l'\\ 


■('ill 


i'    I 


t    i 


roady  nu'iitioiifd.  Fur  fxamplf.  if  tlu'  animal  is  hold  up  with  the  hiiul 
loRs  hnntrinir  down,  thi<sr  will  (>l"lcn  exhibit  rhylhniip  floxion  nnd  Pxton- 
sinn  iniivoniPiits.  with  tlii'  two  liiiilis  ai'tiiiR  altfinntcly,  ns  thoy  would 
in  wnlkinc  or  runninir.  This  is  soiiK-timos  cnll'Ml  the  mark-time  rfflf.r. 
Another  eonii>liented  movement  may  he  produeed  hy  plneinR  the  nnimal 
in  wafer,  when  it  may  make  the  movements  of  swimminp,  hut  its  swim- 
min-.'  will  not  he  snffleient  to  keep  it  on  thr  surfaee.  These  swimminir 
movements  afe  more  perfeet  in  the  spinal  frog. 

After  eomplete  reeovery  from  si)inal  shock,  the  hind  linihs  are  more 
or  less  in  a  eondition  of  rrtniKitu)  rnnlrnrlurr:  the  vnspulnr  nnd  other 
\  iseernl  reflexes  are  in  perfect  eondition,  nnd  a  tnnrkcd  rise  in  hlood 
pressure  oeeurs  when  one  of  the  sensory  nerves  of  the  hind  liml)  is 
stimulated— nn  experiment  whieh  enn  he  performed  in  sueh  nnimnls 
without  the  administration  of  any  anesthetie,  sinoe  the  nnimnl  feels 
no  pnin.  In  female  spinal  animals  imi)re(ination  niny  oeeur  nnd  preg- 
nnney  proceed  in  normal  fashion  neeompanied  hy  the  usual  seeretion 
of  milk.    The  siirnifieanee  nf  this  faet  will  he  dwelt  upon  Inter. 

SPINAL  SHOCK  IN  MAN 

As  we  ascend  the  animal  scale  we  find  that  reeovery  from  spinnl  shock 
fakes  lonirei-  and  lonsrei-  to  occiir  and  hecomes  less  nnd  less  perfeet.  Tn 
fh'  case  of  man,  recovery  is  never  complete,  foe  a  permanent  eondition. 
vlnch  has  licen  called  "isolation  dystrophy."  supervenes  hefore  the 
symptoms  of  shock  have  lieeii  recovered  fmni.  The  tendon  .ierks  are 
jiermanently  aholished  in  complete  lesions  of  the  cord  in  mnn,  nnd  even 
when  the  lesion  involves  only  one  lateral  half  of  the  cord,  this  reflex 
is  either  entirely  absent  or  very  feehle  on  the  eorrespoudinp  side,  thouffh 
normal  on  the  other  f Holmes' V  Severe  lesions  above  the  lower  dorsal 
refion  practically  always  leave  the  lejrs  in  a  permanently  flaccid  con- 
ilition,  with  aceom)),inyinf.'  atro])l!y,  but  sometimes  automatic  movements 
of  flexion  and  extension,  like  those  of  the  mark-time  reflex,  mny  set  in. 

When  the  injury  of  the  cord  is  less  severe,  the  limb  musculature  is 
llaccid  and  toneless  for  some  time,  the  tendon  .icrk  and  the  abdominnl 
and  cremasteric  skin  reflexes  bcinjj  entirely  absent.  After  some  time, 
however— it  may  be  as  early  as  ten  days— the  muscles  begin  to  reac- 
f|uirc  some  tone,  and  a  little  later  the  tendon  .ierk  becomes  elicitable. 
Hejrardinjr  the  b<havior  of  the  flexion  reflex  after  sjnnal  in.iurios  in 
man,  it  has  been  fuiinil  that  tlu'  part  of  it  known  as  the  Hakinski  reflex 
is  not  elicitable  after  severe  lesions,  but  in  those  that  are  less  severe 
a  flexion  of  the  (jrent  toe  may  occur  on  stimulation  of  the  sole.  Later 
this  movement  may  be  accompanied  by  contraetion  of  the  hamstrings, 
and  later  still,  in  favoiable  cases,  by  Mexion  at  knee  nnd  hip.     Tn  these 


i«i.\ 


KIl'I.IXKS    IN    Tin.   hnVAl,    AMMAI-    ANI>   KriNAI.   KIIO.  K 


807 


,,,s..s  also  tlio  Unl.inski  nllox  chni-K.-s  fn.m  n  fl.'xion  to  nn  extension 
„f  the  «re«t  foe.  It  is  important  to  note  in  eonnertion  with  the  nbovo 
association  of  n.ov.me..ts.  that  the  s.M.so.y  area  of  the  sole  is  eonnecte.l 
„i,h  the  same  s.r.nent  of  th-  M^ioM  -onl  tlii.t  furnishe.  ♦»";""'<•«• 
fil.ors  to  the  flexors  of  the  to.-s  nn.l  the  han.stri..,rs  (first  snernU     The 

,„.ov,.r.v  after  si k   therefore  sets  in   earlier   for   unisefftnental    reflex 

Mr.  lis  than  for  thos.-  iiivolvioL'  several  seirinents. 

The  Cause  of  Spinal  Shock 

The  relaiionship  of  th.  ,.rofn„.litv  of  spinal  shoek  to  the  phylo«enetie 
position  of  the  animal  in.lieates  that  the  shoek  must  he  .Ine  to  the 
isolation  of  the  lower  spi.ial  so.Mnents  from  the  ln<rher  renters  fP.ke  _V 

I,  ,,ns  heen  sue te.1  tl.at  the  spinal  seetion  in  the  hiirher  bnt  not  in 

the  lower  animals  breaks  a  nervons  pathway  in  wbieh  normally  the 
,.rflev  innn.lses  travel.  Aeeor.linc  to  this  view,  the  afTerent  .mpnlse 
,vhen  it  eutevs  th.  s„innl  eor.l  in  the  lowrr  nnin.als.  ehooses  the  shortest 
possible  ronte  to  the  efl-eetor  nenro,  of  the  same  or  elosely  a^lli.eent  seff- 
,„onts  bv  the  eollnteral  brnnehes  snrinirintr  from  tbe  srnsory  neuro,^ 
Tn  the  higher  animals,  bowever.  it  wonl.1  appear  that,  althoncrh  this  loea 
spinal  patbwav  is  present  an.1  mav  be  taken,  yet  it  is  nsnally  passed 
bv  an.l  the  impulse  travels  up  to  tbe  higher  eenters,  from  wh.eh  it  is 
,lH.n  trnPsn,itte.1  bv  the  pvramirlal  traets  to  the  motor  neurons  eon- 
,.orne.1  This  wonl.l  appear  to  be  tbe  patbwav  for  nervons  reflex  im- 
„„ls..s  m  bi.'ber  nnimab  the  beaten  traek.  When  the  spn,al_  eord  is 
s.vere,!  therefore,  tbe  eonrlition  of  sboek  supervenes  beenuse  impulses 
,,„ve  ,u'.t  vet  learne,l  that  thev  may  fi.ul  a  shorter  ror.d  to  the  motor 
neuron  bv  tbe  eollateral  than  by  the  patbwav  wbieh  they  usuallv  travel. 
Thev  barn  this  only  after  some  time,  wbieh  explains  the  slow  re- 
eoverv  of  the  reflexes  from  sboek. 

Tt  is  obviouslv  a  .liffieult  matter  to  supply  -lireet  proof  in  support  of 
„„.  „l„.ve  bvpothesis  of  tb-  eause  ..f  spinal  sho-k.  but  besides _ the  ,ti- 
.lireet  evi.lenee  furnisbe.l  bv  observations  on  the  de^^ree  to  wb,eh  this 
..on.lition  supervenes  in  .lifferent  groups  of  anmials.  the  hypothests 
nlso  eonforms  well  with  all  tbe  other  faets  wbieh  we  know  resardms 
„,„  ,..„„ii,i..„.  For  example,  it  is  well  known  that  tbe  portion  of  the 
,,o,lv  above  the  transeetion  of  the  spinal  eord  in  no  way  suffers  from 
the  shoek  i^herrinston  has  .leseribe.l  a  monkey  the  eord  of  wh.eh  was 
out  below  the  eervieal  reiri-.n.  and  wbieh  immediately  after  the  opera- 
tion  amused  itself  by  eatehin?  flies  with  tbe  anterior  extremities,  whereas 
the  posterior  extremities  were  in  a  eouditior,  of  the  profoundest  shoek. 
Sueh  experim.uts  further  indieate  that  the  shoek  ean  not  be  dependent 


!     ;   s: 


2i 


■!  '  r'lJ 


i  IrM 


808 


Till. 


INTHAI.    NKHVOIH   KVKTKM 


upon  the  lowi'iiiitf  iif  nrtci-ial  IiIimkI  prcHMure  wh'u-h  n  Nrrtlon  of  tho 
iMiril  liinlu'i'  til  .11  till-  iniil-iliiiHiil  rcifioii  must  fiitail.  Tho  poor  mitritivc 
condition  of  tho  hIum  whii-h  we  havo  «'i'i>  to  ixist  in  the  liiiul  limlm 
In  shock,  tihowH  that  the  lilooil  vessels  in  them  arc  iiiofouiully  dilated, 
hut  evidently  the  fall  in  hlood  pressure  has  iiothintr  to  tlo  with  the 
faulty  (iinduetii.ii  Ihrouirh  the  spinal  eord.  for  ^ueh  a  fall  wmild  affect 
the  centers  for  the  fine  limits  as  well  as  those  for  the  hind,  and  yet 
the  former  show  no  sym)itoms  of  shock. 

Exactly  similar  slicek  is  ohtaiiied  liy  any  section  of  the  spinal  cord 
as  hijrh  up  as  the  medulla.  Of  course  as  tiic  sect  inn  is  made  hitfher  and 
hijrher  up,  the  rcsultinc  i)aial\sis  heconu's  aiore  and  more  marked,  nnd 
niny  reach  such  a  decree  of  severity  that  ri very  of  the  animni  be- 
comes an  impossihiity. 

When  we  come  to  c.nsider  the  functions  of  the  various  parts  of  the 

hrnin,  we  shall  have  ( asion  to  study  the  elTeets  following  section  at 

hiifher  h-vels  of  the  cerclirospinnl  axis.  Meanwhile,  however,  it  is  im- 
portant to  note  that  when  a  section  is  iiuitic  across  the  crura  cerehri,  so 
that  the  eerehral  hemispheres  alone  are  isolated  from  the  rest  of  the 
nervous  system,  a  condition  of  contracture  of  all  of  the  extensor  muscles 
occurs.    This  eonditi m  is  known  as  ilt<<rt„riil(   riijiilitii. 


1 

i 

■  i 

1  ,1 

1^   :, 

1 

ill  '' 

f 

III 

i!  i 

; 

' 

il 

l.s 

■il 

CIIAI'TKH  X«' 
PHYSI(»r,(Hil('.\I-  I'KOI'KKTIKS  (»!•'  TIIK  SIMIM.K  UKFI.KX  AUC 

\V..  ninv  now  ,.. .M.l  t..  slu.ly  tl.o  i.n.p,.iti..^  of  .vflox  n.'fion  orrur- 

rinir  fl.ioimli  tl...  iM.lnfr.I  H|,innl  .Tnt.TH  of  n  spinnl  animal.     TIhtp  nrr 

two  HNiH-rts  of  tlu .•stion  to  l.o niMrml:     (n   Ww  i.ro,,..rtios  of  n 

sinifh'  rvnvK  an-,  „n.l  i'D  tl..-  aHlo,,  or  inllu.-n.-o  of  on.-  rc-Hox  arc  on 
nMolhor  Til.-  in.i...itan.-,.  of  th-  laft.r  will  1..-  «-vi,l.-nt  mLom  it  is  rc- 
nion.l.rrr.l  that  .-on.pli.-af.-.l  iniiH.-iilar  nmvc-ni.-nts  .l-pon.l  for  thnr  proper 
coordinntion  rntir.-lv  on  th.-  inf.-ra.-tion  l..-Uv..f-n  tl.o  various  r-d.-x  arcs 
which  <-onipoH,.  11,..  n.rNoUH  sysL-n..  This  int.-raotion,  as  alnn.ly  rx- 
plninr.l,  has  I.......  rnll.-.1  l.y  Sh.-i-rin^ton  tlu-  int,,,n,lio»  of  the  urrrouf 

»\l»trm.  . 

Pn.lmMv  th..  sin.pl.-st  way  to  stu.ly  tl...  physioloir.o  proportios  of 
the  simpi..  r.-n.-N  is  to  .-on.pan-  th.-  m.o.I.-  of  ron.l.iot.on  ..f  n 
lUTVo  impiils.-  thronu'l.  it  with  .-..n.liK-tion  alonif  a  sin.pl.>  nrrvc  trunk. 
By  romp- rinif  tho  two  n.o.l.-s  ..f  .M.n.lu.-tion  wr  shall  ho  h.'ttor  ahlo  to 
npprooi/.to  tl...  n.o.lifi<-ntions  to  wl.i.-h  th.-  in.p.ilso  is  s..l.,i..ot<..l  hy  con- 
duoti..n  thronjjh  tl...  r..fl.-x  ar...     The  ii..p..rtant  points  am  th..so: 

1  The  Latent  Period.  Th.-  lat.-nt  p..rio.l.  or  p.-ri...!  whi..h  intrrvTiiPs  he- 
twpon  tho  niomont  of  applh-ation  ..f  tho  stin.uiiis  an.l  tho  rosp.inso.  is 
vrry  sh..rt  in  tho  oaso  of  a  n.-rvo  t,-iinl<.  an.l  un-lor  n..rinai  oon.lit.ons 
nlwavs  tho  san.o.  hut  is  .piito  va.ial.l.-  an.l  s.nn..tii....s  vory  lon«  in  tho 
«n«o  of  n  redox  an-.  Thus,  in  tho  oaso  of  tho  oonjiinotival  rollox.  whioh 
is  pr.«l.i..o.1  hv  ai.ph  inif  a  stinn.lns  to  tho  o..r..oal  oo..j,..notiva  (oaus....-' 
a  ol<.sin«  ..f  tl.o  ..v.li.ls),  tho  nHox  tin...  is  vory  sln.rt  an.l  invanal.lo. 
whoroas  in  tho  oaso  of  tl.o  s.-ratoh  rotl..x  it  may  vary  from  two  an.l  a 

half  to  thr..o  nn.l  a  half  s.-.-oiuls,  a. nlinn  t..  tho  stromth  of  tho  st.ir.u- 

lus.    The  s..at  ..f  .lolay  i..  th.-  rollox  aro  is  p..<.l.al.ly  in  tho  synapse,  hut 
its  .■i..is(.  is  ohscurc. 

«>  Grading  of  Intengity.--!..  a  i.orvo  trunk  th.-  intensity  of  the  un- 
puNo  is  n....'o  ..r  l.-ss  prop..rtional  to  tho  st.vnjrth  ..f  tho  stimulus.  This 
ean  he  ju.k'o.l  hv  ol.servinu  o.thor  th.-  n.-tion  our.T..t  i..  tho  uorte  hy 
moans  ..V  a  }ralvnnon..-tor  ..r  tl.o  .•.■sp.n.so  of  tho  .-...l  o.-sxan;  o.  !.'.,  n.usole. 
„,ta..l....l  t..  11..-  ..orvo.  1.,  tl.o  .-ase  ..f  a  r.-H.-x  aro,  on  the  other  han.l 
there  is  hv  i...  n.eans  s..  ovi.lout  a  pa.all.-lis.n  hotwoon  stimulus  ami 
r.-sp..nse.  ■h-IIoxos.  h..wover.  •  ..ry  .•onsi.lo.ahly  in  this  roirar.l.  The 
..onjuMolival  n-ri.-x  an.l  tho  .-xt.-nsor  thrust  hehav..  a.-.-o.-.l.....'  to  the  so-.-alh-.l 
"all  or  n..thin..'  prinoiplo:"  i.e..  tl.o  i..to.,sity  o(  tho  r.-sponso  is  more  or 
loss  in.lcpon.hnl  of  tho  stron-th  ..f  tho  stimulus  I"  ..thor  rotlox.-s.  suoh 
as  tho  lloxi..n  rollox  an.l  tho  s.-.atoh  .•o(l..x,  tl...  i..t..nsity  of  tho  r..sponso 

800 


.! 


li 


i  !   i 

'       lii 


•;  it: 
i.; 


li 


ikftM-^'ili' 


i^:i 


'■I  i 


,  s 


M() 


IHE   (  KNTHAI.    NKRV()t:s   SYSTKM 


is  iinicli  more  noaily  proportional  to  the  stronRth  of  the  stimulus.  Thus, 
a  fpnblo  stimulus  applied  to  the  flank  calls  forth  only  a  slijibt  flexion 
of  the  hind  limb  of  the  same  side,  whereas  a  stronger  stimulus  sets 
(rninp;  a  typioal  seratehinR  movement. 

n.  After-eflfect.-  When  a  stimulus  is  removed  from  a  nerve,  the  effeet 
whieh  it  produees,  as  Judsred.  for  example,  by  the  netion  current,  im- 
tii.'iliatcly  disappears.  TbiTi-  is  no  after-resixmse.  In  reflex  ares,  how- 
ever, sueh  a  phenomenon  is  usually  well  marked.  I'artieularly  is  Ibis 
the  ease  in  the  flexion  and  serat.'b  reflexes  of  the  si)inal  doff.  A  mo- 
mentary stimulus  of  o]>tinial  strenu'tb  applied  to  the  scratch  skin-area 
may  produce  no  immediate  response,  but  after  its  removal  a  violent 
scrat.'bin-r  movement  may  set  in.  This  after-discbar!.'(\  in  cases  in  which 
tlie  stimulus  is  stnmtr.  may  indeed,  as  in  the  flexion  reflex,  be  more 
inaikiil  than  the  response  duriusr  the  time  of  application  of  the  stimulus. 
In  this  particidar  reflex,  the  after-discharge  often  lakes  the  form  of  a 
cloinis,  with  a  rate  of  contrac', ,011  of  from  seven  and  a  half  to  twelve 
per  second.  The  crossed  extension  reflex  also  Iuin  a  very  pronounced 
nffer-discbarfie,  which  may  outlast  the  stimulus  for  from  ten  to  fifteen 
seconds.  Kcirardinjr  the  phenomenon  of  after-discbarfre.  Sherrington 
has  stated  that  there  is  "no  feature  of  the  conduction  of  a  reflex  arc 
w  ieb  distinsruisbes  its  mechanism  more  universally  from  that  of  a 
neivc  fiber,  tract  or  trunk  than  leiisrlhy  after-discharsre." 

4.  Summation.-  AV ben  a  subliminal  stinudus  that  is,  one  that  has 
in  itself  no  visible  efTi-i-t  is  frcpiently  reiteated  in  the  case  of  a  nerve, 
no  response  occurs.  Tn  the  case  of  a  reflex  arc.  however,  such  repeti- 
tion of  subliminal  stimuli  ultimately  calls  forth  response.  This  sum- 
mation is  very  evident  in  the  case  of  the  scratch  reflex:  \  s,  one  or 
two  ele.-trical  stimuli  applied  to  the  scratch  field-area  call  forth,  as  a 
rule,  no  nn.vcMient  of  the  corresponding  bind  le<r,  but  if  these  same 
stimuli  ar.'  frequently  -eiieated.  the  typical  reflex  scrat(diin<,'  movement 
will  oeiMU'.  Evidently,  then,  in  a  reflex  arc  there  is  ;i  considerable 
amount  of  resistance  towards  a  siu'.'lc  stimulus,  which  resistance  is 
ovcreoMie  by  a  sueeession  of  stimuli.  Tn  other  words,  the  threshold  of 
the  eScit.Mbility  <if  the  reflex  nu'cbanism  becomes  lowered  as  a  result 
of  its  pievious  stinndation.  Kaidi  stimulus  excites  the  sensory  surface 
so  that   it   n'spoiids  more  easily  I0  the  sueeeedinir  stimulus. 

.'  Irreversibility  of  the  Direction  of  Conduction.  This  is  well  illus 
trated  in  the  so-ealled  r.eli-Mai.'eMdie  law  of  e(uiduction  in  the  spinal 
ner\-e  roots.  A  motiu'  impulse  travels  out  of  the  eord  by  the  anterior 
roots,  .vbile  ;i  sensory  imi>ulse  travels  in  by  the  posterior.  This  direc- 
tive influence  can  uol  depend  on  the  nerve  trunks  or  'he  nerve  cells,  for 
nerve  trunks  conduct  equally  in  both  directions,  ami  so  also  nnist  the 
nerve  coll.     The   irreversibility    must    therefore   clepend   on    the  synaptic' 


I-HYSlOIXXilCAI,   I-KOI'KKTIDS   OK    TUK   SIMI-Li:    KKKLLX    AUf 


Sll 


..onnections.  It  oan  l.e  .lemonst.ated  by  ol.s.rving  tlu-  act....  cur- 
rent produeod  in  th.-  spinal  .•or.l  by  stimulating  the  anterior  nr  posten..r 
spinal  roots.  In  the  former  case  no  action  current  is  observed,  but  it  is 
vorv  evident  in  the  latter  case.  „      ,  ,      c       •      .   . 

,;■  The  Refractory  Period.-This  has  bee,  well  defined  by  Sherrington 
as  beint?  "a  state  durinfr  ^vhich  apart  from  fati.,ue  the  mechanism  shows 
less  than  its  full  excitability.-     We  are  already  familiar  .vith  the  re- 
fvaetorv  period  in  the  cases  of  the  heart  muscle  ami  the  museulature  of 
the  esophaun.s  and  intestine.     F.n-  example,  the  application  of  a  stimu- 
Ins  to  the  .piicscent  fro.  heart  while  it  is  contraetins  in  response  t,.  an  im- 
,„ediatelv  precediiiR  stimulus  fails  to  produce  any  further  effect.    The  re- 
fractory" period  is  extremely  brief  rone  thousandth   of  a  second^    in  a 
nerve  trunl<.  but  is  much  lon.er  in  a  reflex  arc.  beina  probably  lo.,gest 
in  the  case  of  the  serat.d,  reflex,  in  which  it  is  demonstrated  by  the 
fact  that,  however  fre.,uent1y  we  apply  suitable  stimuli  to  the  sensory 
surface    the  rhvthm  of  response  of  the  contractinfr  limb  is  always  the 
same      After  each  stimulus,  therefore,  a  refractory  period  must  become 
deveb.ped  durinfr  which  a  repetition  of  the  stimulus  has  no  elTect.     It 
is   evident    that    the   existence    of   the    refractory    period    is   the    factor 
responsible  for  the  rhythm  of  the  movements. 

Tt  is  intcrcstin.r  to  consider  the  exact  structure  of  the  reflex  arc  that 
is  responsible  f..r  the  existence  ..f  the  refractory  ,,hase.  It  obviouslv 
,;,n  not  be  a  function  of  the  motor  neuron,  for  through  the  same  motor 
neuron  mav  be  diseharped-  at  one  time,  impulses  which  brin.  about  the 
seratchin..'  movement  and,  at  anoth..-.  those  causn,..-  a  tonic  flexion  of 
the  same  muscles.  Nor  can  the  seat  of  the  refractory  period  be  in  the 
sensorv  area  of  the  skin  or  the  afferent  murou.  for  if  a  s,.ratch  move- 
,neiit  is  elicited  bv  stimulation  at  a  point  .1  in  the  proper  skm  area. 
1,,c  rhvthm  of  response  which  i.  calls  forth  will  not  ,n  any 
wav  be  altered  bv   the  application  -f  a   secon.l   stnnulns  a,.p bed   a     /> 

Mt  some  distance  from  .1   and  havi,...'  a  .litTcrent    f> neney   (In.'.  -11'. 

There  is  evidentlv.  therefore,  some  part  of  th.  reflex  are  that  .s  eommoii  to 
in.pnlses  startinu'  botl,  at  .1  and  at  /.'.  for  if  in  each  of  these  spots  a  refrac- 
,.,rv  phase  oe.M.rred.  then  there  would  Ve  interferen.-e  b.-foiv  the  two  im- 
nnlses  had  reached  the  eent<.rs  of  the  spmal  .ord.  T^y  exclusion,  there- 
fore "the  seat  of  the  refraetorv  phase  seems  to  lie  s.m.ewhere  central 
,„  th'e  reeeptive  neuron  in  the  afferent  are"'     f Sherrington- V 

Manv  other  types  of  reflex  activity  illustrate  rhythm  due  to  tlu^r."^ 
fraetoiw  phase.  Two  laboratory  ..xamples  may  be  «iven:  (V  men 
the  central  end  of  an  afferent  root  is  stimulated  m  the  lumbar  region 
of  ,l,p  spinal  cord,  the  movement  produced  is  distinctly  rhythmic  m 
eharacter  f2>  Tpon  stimulatinir  the  eentral  en.l  of  the  sciatic  nerve 
in  a  fropwhose  spinal  cord  has  been  cut  some  davs  previously,  a  clonic 


': 


t! 

rl 


J 


l> 


n,  ilM  -V 


S12 


Tin;  (KNTii.M.  Minors  systkm 


fiction  of  the  ('(mlraliilfriil   foot  occurs,  aiitl  the  rate  of  the  rhythm  is 
not  alTected  hv  variation  in  the  fre(|ueney  of  the  stimulus. 

Ill  all  the  ahove  cases  the  refractorv  period  iiinv  be  heltl  responsible 
for  the  rhythmic  nature  of  the  contraction.  In  other  reflexes  it  e.\ists 
for  anotlier  i)iirpose.  In  the  case  of  the  extensor  thrust,  which  it  will 
be  rememberc<l  is  elicited  by  pressure  applied  to  the  i)ads  of  the  jdantar 
aspect  of  the  foot,  the  momentary  extension  o{  the  leu  lasts  only  for  .'» 
little   less  than   two-tenths  of  a  .second,  but   is   followed   by  a   refractory 


u 


aM 


,J^' 


iJ 


ll'l 


lis^    !  ifitl:''' 


nil* 


III  1  i 


!.     1 

41  J 


).■,„  'II.  I'r.uini:  tr.im  tlu  Inii.l  I'liiti  "f  .1  ^l•ln.■ll  .1"|;  .liiriil!  tlu  s,  r:il,  liinj;  lu.iv.  in,  iits  pi" 
ilm>-.l  l.v  ai.|.l\MH!  -imuili  at  I«"  >kin  pciiits  (.1  ami  11),  Ih.'  ;,|.|.li.ali(.ii  of  llir  -tiiiuih  l.riiiK  iii 
•  luatiil  liv  tin-  viK^aN.  N.  t  i.iilv  \n-n-  llii'  'timiili  .iii|'tiiii  .11  ililTiriiil  ihmiiIs.  IhiI  al  /I  liny 
«.!,•  "f  imuli  kT.ali  r  IT.M'Hn.i  tlan  at  .1.  .Mtli.'iiiih  then-  !■.  a  •■\\ghx  i  Ii.tiiki-  in  "Icnal  sixii."  it 
Hill  hr  <il.-,ivii|  that  I'j.ic'  I-  ii.i  altiration  in  rlnthiii.  iii.lii  almi!  that  llii-  piuimly  niu-t  I'l-  a 
fniKti'.ii    i.t   ;li''   I'liai   ii.i-;i!ii.ii   i..tli.      (  I'roin   Sh,  ri  inKt"n.) 

period  lastiiii;  nearly  a  whole  second,  duriiiir  which  a  second  stimulus 
elicits  no  response.  The  oli.jcct  of  this  Ions;  refractory  |)eriod  is  no  doubt 
that  oi)portuiiity  may  be  tiiveti  for  the  tiexor  muscles  to  perform  the 
contraction  that  woiilil  iialurally  ensue  tlurin^'  the  normal  occurrence 
of  tlic  extensor  flniist,  as  in  the  act  "f  walkiii}":.  When  the  .'inimal 
places  his  foot  on  the  "iidiind,  the  sutbleii  |M'essure  exerted  on  the  iiatl 
of  the   foot    immediatciv   calls   forth   the   extensor  thru.st,   by   means  of 


PHYsioLOdicAi,  I'uon  KiiKs  <.F  Tin;  snii-i.i:  hkki,i:x  Aur 


813 


^vhi^h  tl.o  ^v.M«l.t  .(  tl.o  1..m1.v  is  t,..ni...nnil.v  rcm'.voa  from  ll.o  Rroun.l, 
ami  the  nmsHcs  p(.rf..nn  llu-  ,.ont.a.-tio„s  n.-.-i.ssnry  1„  pnulvu-o  iloxu.n 
of  .1...  H.nb.    Alt.,o...l.  11>,>  n.f,a..t.n,-  po.io.l  is  un.lT....t...l  l.y  t ho  s.n-n^  1 
of  tl.o  stimulus  it  is  v.M.y  .l.M..'"<ln.t  upon  11,-  intornn    .•on.l.tu.n  of  to 
„..no  roflox  ar...  su.-l.  as  tl.at  ..auso,!  l.y  .■han.^es  i..  l.loo.l  supply  or  In 

'''nri!''r.on,h,rli<n,  i. /'  I'ss  nsislnnt  than  ...tvo  ..o.ulu-tion  to  various 

.oM.litions  alT..otin«  tl.o  nufitivo  oomlition  of  tl.o  oon.luot.uR  pa1l.^vn^■. 
For  oxa..,plo.  .lop.ivation  of  oxy.ou  oausos  l.ut  sli.ht  ''^^-^'-'-'-'J';^; 
tho  oon.luotion  alo,.,^  a  uorvo  tnn.k.  but  vo.^y  soou  a l.ol.shos  the  p.nal 
rolloxos  Kvou  iu  tho  tVoj,'.  .■•■Hox  u.ovomonts  outiroly  d.sMppoar  in  th.r  y 
to  fortv-fivo  ...iuutos  aftor  tho  oouto.s  l.avo  l.oon  .ou.lo.o.l  oou.plotoly 
anon.io:  a...l  in  n.au.n.als  thoy  -lisappoa,'  i..  a  fow  n.inutos  In  the  ease 
of  .lr,..'s  suoh  as  ..hh-rofonn.  <».:i  por  .■•■nt  of  tho  ,lru.'  ...ay  bo  ro.,....o.l  to 

aholish  oo.Mlu.-tion  i,.  a  norvo.  whovoas  a  n.,.oh  lowo.^  po.rontairo  ,s  suO,- 

<•io.it  to  aliolish  it  in  a  roflox  arc. 

From  tho  al.ov..  .lifro.c.oos  in  oo.,.l..otion  in  a  norvo  <'""  <  ""^  ;•■; 

fl..x  a..',  uo  loa...  ma.,y  faots  oon,.o...i..«  tho  iu-portan.-o  of  tho    atto,. 

ana   ^vo  f,..'tl.or  s.o  that   tho  .lifTorono.-s  a.o  .luo   vo.y   lar^oly  to  the 

synaptic  coniioi-ti"... 

SUCCESSIVE  DEGENERATION 

r...for. ..oh.-li...'  tho  suhjoot,  it  n.ny  ho  of  i..to,.ost  to  oonsi.lor  h.'iofly 

„,,  „,.,„o.l  of  snnr.s;rr  .hyrnrration.  hy  wl.ioh  Shorrin-.ton  snoooo.lo. 
in  .lomonst,ati..u'  tho  oxaot  traots  in  tho  ^vl.ito  mattor  of  tho  sp.nal  oo,.l 
alons:  xvl.ioh  tho  intn.spinal  ,.ou.o..s  l.'avol  f.nn.  ono  sovMno..1  to  a,.otho.. 
This  was  .....ko.!  out  i..  tho  ,.aso  of  tho  son.fh  rollox  ,n  tho  follow..,!,' 
man..or:  Tho  spinal  oo..,l  was  li.st  of  all  o.,t  in  tho  upi---  "7'";";;7'2 
s,>  that  ,lo,..,.o..ation  o.-on.To.l  i..  all  tho  .losoo.„ln,u  t.a.-ls  holou  tl.o 
lovol  of  tho  scotio...  I.,  about  a  yoar's  ti.no  those  ao^.o.iorato.l  t.'ao  s  ha.l 
on,in.ly  .lisappoa,.,!.  a.,.l  .ho  -lobris  of  th.;  do.ono.a.o.l  fibo.s  ha^  boon 
,,,,a,.o.l  bv  ..i,-at.ioial  tissuo,  so  that  a  soot.on  o.  tho  oo,-.  .•ovoalo.l  noth- 
ing but   h-'-althv  novvous  tissuo  with   oioatrioos   who.o  tho  ,lo«o.,o,.ato.l 

t,;,.ts  ha.l  oxis"to.l.     Who.,  at  this  stasro  a  s .n.l  .'ut  was  ma.lo  aoross 

tho  <..n-.l  a  littlo  lowor  tha..  1ho  ti.'st  ono.  furthor  -b-u'onorat.on  ooourro.l 
involvin..  thos..  (ibo.s  whoso  .•o.,to..s  wc.r  b-o.to.l  botwoon  tho  two  ouls- 
th-.t  is  tho  1ibo.N  oon.b.sr  f.'o.n  tho  intrasi.b.al  no...-o..s,  w.th  tho  oohs  ot 
whioh  'tho  a(To.T..t  ..orvo  tibors  oou.i.-s  fnn..  tho  skin  of  tho  soratoh  ro- 
tlex  a.Ta  wo.-o  .-o.n.o.-to.l.  A  so.-tio..  of  tho  oo,.l,  stab.o.l  app.'opr.atoly 
for  .lo.'o..orato.l  f.bo.^s.  at  this  tiu.o  ao..,onst,ato.l  thoso  fibo.'s  to  ox.st 
in  tl.o  rato,.al  oob.n...  of  whito  n.atlor,  ih.is..  that  t.avol  a  short  (l.stnncc- 
i.c.  hotwoon  noi-hbori..}-'  soi,Muo..ts-~boi..jr  noar  tho  siray  ...atto.-,  and 
those  ti-avoliiiK  (x.-oator  distauoos,  towai-ils  tho  outs.dc. 


•     h 


ill 


! 

1 

.  i 

•;h'; 

^•1  hi   nil 


:ii    li 


):      ■  1 


•i      ! 


rllAITKH  .\(  I 

RKCIIM{(K\Vr-  INNKUVATION 

Reciprocal  Inhibition.  It  murlit  iii)pt'nr  tluit  to  in'iul  a  juiiit  or  to 
nicivc  tlic  ryolinll  tlif  only  iimsciilnr  action  vciniircd  wonM  he  fontrac 
tion  of  tlio  iiuisclos  wliicli  flox  tlio  .joint  or  rotate  tlio  oycball,  nnd  that 
tlip  antajronistic  iiiusclcs  would  niorcly  liecomo  ]>nssivc1y  rlntiijated. 
Wlicn  wo  rcnicinlicr,  however,  that  all  the  nuiseles  of  the  hody  are  or- 
dinarily in  ; idition  of  sliirht  eoiiti'a<'tion,  or  tone,  and  that  this  tonds 

to  hepomo  iiierensrd  when  the  museles  are  passively  strotehod.  then  ^^  e 
see  that  for  etiicient  movement  there  must  he  inliiliition  of  the  tone  oT 
the  nuiseles  whieli  o]ii)oso  those  that  are  eontraotinjr.  This  rfciprocnl 
iiililhitimi.  as  it  is  ealled.  is  a  very  widespread  fnnetion  thronshnut  the 
animal  body.  Sometimes  it  is  purely  pei'iphoral  in  oriffin,  ns  in  the  elaw 
of  t'  M\tisli.  wiii'ii'  stiiiinlation  of  the  ner\e  causes  an  oi)einii}r  of  the 
.  elav*  V  to  the  eontraetion  nf  one  set  of  mnseles  and  the  simultaneous 
inhiliiiinii  (if  tlieir  antatroiiists.  Tnstanees  of  prn'i>lifral  rcciprornJ  in- 
hihitit)!!  ill  the  hiu'hei'  animals  are  not  so  common,  hut  are  illustrated  in 
the  ease  of  the  myenterie  reflex,  where  it  will  he  rememhered  a  eontraetion  nf 
the  intestine  ovi-r  a  holns  of  food  is  accomi>aiiiel  by  itdiiliition  in  front  of 
the  bolus.  The  reei)iroeal  action  in  this  case  is  probably  dependent  on 
the  myeii'eric  i)lexus. 

On  the  other  hand,  n cifirociil  hihihiiion  nf  cnilrnl  nrlfiln  is  very  com- 
mon in  the  hiirher  mammalia.  Thus,  in  the  case  of  the  lateral  movement 
of  the  eyes,  if  we  cut  the  third  and  fourth  nerves  to  one  eye.  sav,  the 
left,  the  external  I'eetns  of  that  eye  will  alone  be  under  the  control 
of  the  ner\ous  system,  throuirh  the  sixth  nei've:  nevei-theless,  if  we  after- 
^^■ard  cause  the  animal  to  lool<  vard  the  I'iirht,  as  by  holding  some  ob- 
.ject  in  that  <lirection,  it  wU  .md  that  the  left  eye  as  well  as  the 

rii.'ht  follows  the  ob.jeet.  Obviously  there  must  be  an  inhibition  of  the 
external  rectus  muscle  of  the  left  eye.  an  inhibition  which  is  iironounced 
enough  to  briliL'  about  a  movement  of  the  e.xeball,  and  which  exactlv  cor- 
lesponds  in  i>oint  of  time  ^ith  the  contraction  of  the  external  rectus  of 
the  ritrht  eye.  This  nM)vemeut,  due  to  the  atonieity  of  the  external  rec- 
tus, docs  not  however  succeed  in  causin^r  the  eye  to  rotate  beyond  the 
midline  of  the  field  of  vision.  This  is  an  instance  of  a  wiled  reeiproeal 
inhibition  ;  i.  e.,  a  reciprocal  inhibition  brouffht  about  by  stimuli  eominsr 

814 


KKCII'KIM  Al.    INNl.KS  \TlciS 


Hi:. 


,V.,m  the  voliti....al  .■.•ut.r  in  tlu-  .•er.lnu.u.     The  same  .•m.lt  nmy  l..« 
!l,rinccl  by  ..loetri.-  stinn.lation  of  the  ..enter  for  eye  moven.ents  on  the 

""Tir!uosrim,...rtant  details  eonee.ninK  the  .ueehnnis.u  of  reeiprocal 
innervation  have  l.oen  obtained  by  stuayin-,  the  iloxion  reflex  .n  a  spurn 
;;;  •l.lar.hich  has  completely  recovered  fr.un  shooU.  I-uch  an  annn. 
,ho  tonus  of  the  extensor  nu.seles  of  the  knees  .s  well  marked  Th. 
,onus  is  maintained  by  afferent  in.p"lses  transmitted  to  the  spn,al  cord 
r  eceptors  situated  in  the  muscles,  and  its  decree  of  mtens.ty  can 
ho  ostinudod  bv  the  briskness  of  the  knee-jerk,  which,  it  wdl  be  remem- 


1,    .■..„,,,.,  I,-, I    wilh   the    extensor    muscif   nf    ilie    km-c.      lluiiin: 

,.„..    JIJ.      K.o.r.l    t...n    ..•.y.^r.irl;    Vhl   ,1   , ^r     e      Ppos-te    foot   «a.    stinr,,!,tt.-.l.    thus    cans,„« 

,1,,   „„K-  n,arU>.l  l.y   the   lower   »,«,....  the   ^'^^'^^^'..f  Wis    stimulation,    faradu-    >h,uks    were    a,,- 

,he    erossvd    '-'"'-'""    ^'^f;;;;  „^\^^  .^T^     J<<:^'-^'i   ''X   «'-   "PPer   signal..   w,.h    the    rev.,1. 

l,o,od  is  elieitrd  by  tapping  the  patellar  tendon,  and  eonsists  of  .ynd- 
den  extension  ntovement  at  the  knee  .joint.  Hy  observing  the  briskness 
of  the  knee-jerk  we  are  therefore  enablcl  to  form  an  est. ma  e  of  he 
,..„i,itv  of  the  extensor  mnseles;  an,l  if  after  ,loin,  so  we  throw  th.. 
rtfxor;  which  are  their  antagonist,  into  activity  by  clu.t.n.  the  «ex  on 
reflex,  the  knee-jerk  will  be  found  much  less  aetne  we  P™ve"  H 
flexors  from  acting  on  the  knee  joint  and  the  leg  .s  held  -n  an  extended 
position,  irritation  of  the  skin  of  the  le.  will  cause  the  flexion  of  the 


ii 


*  <l 


i 


i 


I    I 


si  ^ 

J  i   ',i 


^  ^! 


n  'I 


!li    ! 


1  ' 

i-i 

! 

i'4!  itv 


><ir. 


Tin:  iiAruM.  nijivois  systi;m 


discoiincctcd  liaiiistiiiitr  iiiusrlcs  sinniltimconsly  with  a  visililc  I'Plnxnfion 
(if  the  cxlcnsois  (  V\<:.  'Jll!).  If  the  let:  is  liclii  prniMTly,  this  n-laxalioii  itiiiv 
lie  niiirkcil  I'lioujrh  to  ciiiiso  a  slijrlit  (Icxion  at  the  Joint ;  and  in  any  onsp, 
if  thi'  kniM'-ji'ik  is  n'<,'iilarly  ciii-itcd  hy  <Miiial  taps  applied  to  tlif  l)ati'lliir 

tend it  will  he  foinid  that,  while  the  llexion   is  i>ein<;  produeed,  the 

knee-jerk  is  very  iiuieh  less  tlian  normal,  if  not  entirely  alisent,  thus  in- 
dieatinj:  that  the  tone  of  the  extensor  iiiuseles  is  diminished.  This  ex- 
jx'iiment  is  very  strikini.'  when  performed  on  a  deeerelnate  animal,  in 
whieh,  as  we  shall  see.  the  extensor  nuiseles  of  the  limli  are  in  a  |)er- 
manent  state  of  hypertonieity  (  i-'i*;.  "21:!*. 

llefore  it  is  perniissihle  to  eonelndo  that  this  reeiprocal  inliihition  is  a 
necessary  event  in  the  movement  of  a  .joint,  we  mnst  iiowever  show  that 
it  oeeiirs  at  exactly  the  same  time  as  the  tlexion  of  the  antagonist.  Rher- 
rinu'ton  has  sncceeded  in  doin^'  this  in  a  considerable  variety  of  experi- 


*^   .Ant. Crural  N. 
[Femoralis) 


I'li;.    ..'1.'.      Iliiti.iiii    -lM«int:    ll 


...111    iNiu  rv.iti.Mi.       I.\fl<i 


nients,  one  of  whieji  we  may  cite  here.  If,  in  a  sjiinal  do<;.  the  tendons 
of  the  llexor  muscles  of  the  knee  .joint  of  one  hind  limh  and  the  ex- 
tensor tendons  of  the  op|insile  lind)  are  cut,  then  the  former  liml)  Avill 
lie  unahle  to  Ilex  jjropeiiy.  hut  will  nevertheless  exhibit  recij)rocal 
inliihition  of  the  intact  extensor  musi'le,  while  the  latter  lindi  will  tlex, 
hut  i-eipiire  i>assi\('  extension  to  Iniiiir  it  hack  to  its  ohl  itosition.  11 
snitahle  siinmli  arc  simultaneously  iippliol  to  the  skin  of  hnth  le^'s  and 
the  movements  of  the  isolateil  muscles  recorded,  the  onset  of  inhiliition 
of  the  intact  extensur  of  the  one  Icu'  anil  the  eonti-action  of  the  flexors 
of  the  o])i)osite  let:  will  he  found  to  airrce  with  regard  to  latent  periods, 
strength  of  rcipiired  stimulus,  summation  and  indeed  all  the  othei-  phys- 

iolo^'ir   •.;'(i|ii'rl  ics  of  rclli'X  .'iCtiiili. 

■Reel,    ocal   iinicrvation  can   also  he  demonstrated  hy  stimulatinsr  'the 
central  end  of  snitahle  afferrnt  nerves- that  is,  eertain  nfTerent  nerves 


Ullil-UdCAI,    INNIKVATION- 


81' 


l-iK.   Jn. 

At  .IK,>.,I  / 
hiljitiiin  "I  t 
(■\t:ilinn  tilt" 
luniii'-al  '"■ 
cii  till-  fl.\. 
l.ilt.r  >lnini 
Ihc   jctiirat 


-Ricinriical    iiimrvaliiiii.      'I  racinu- 


la-  t 


1  a  iliii-nl'v 
...ntraili  '11  '•(  ll"'  tli^"i~ 
.■  acain  llirnwn  iiit"  i"iilr;H' 

i,.ra,a,>.,.    ,..v a;    luivi..    -M    -1,11....    n ;....-.   ;;;^-j'y-,  'X.  'Z 

,„,,i,,.|    :„    ,1,.    saiM.    uir.    .hattlK'    ^::J>:::^:..;:,:^':r  ;;..m„n„iau  -   v>rv 


(vastus    cnirt-iisl, 
11.1111  al:it.tal    lur 

,(  ilu'  i-.ttns..r- 


iiia.U-    t.y    M;y..i:rai  ti-i    I'.i 
:    niti'ili--    (»iiiulin.liiii.~ii- 
al    iRr\i-   w.is   ixiili'I. 
Al  signal   //   lli>'  fl' 


an 


tat. 


:    tlic    l..«ir    MBiial 

saiii.-    line    ll.al   tin'    iM.ii>or«   i-..iilr 

.1       .    ,„„r  ,,■„•   ,,.i".il>.l   ils  iiifliitiui.     Tlii«  .•x|..Tini 
,.,  til.-     ..1 11"  r    .11.    "^  '•;•;,  \        ,,.„„       ,,,,,„„    ShiTringtoii.) 

cc'ltiiuK-t 


111' 

an.l  ill 
■ti.,11  l.v 
i..latiral 

1,   llMll 

.1    til.- 
I.  ally 


ll'i-    n-i'UT.j 


^ 


n<>f 


rt' 


i*  f 


■ 

hi 

1  i'^ 

■i'=i 

I' 

'T  i 

1 

1 

1 

'  i 
1 

i 

i  i 

I 

I  ' 


MS 


III    I  \  \THAi.  Minin 


>Y>TI.M 


nctiiic  1)11  tlio  sniiii'  irroups  of  neurons  will  pioduec  n  (loxion  reflex,  ofheis 
nn  extension  rellex;  thus,  stinnilatioii  of  llie  eoiitralateral  perononl  nerve 
produees  ii  fli'Nion  rellex  oC  the  liiiul  linili  (exeitiitory  for  flexors,  in- 
hihitoiv  lor  exteiisorsi,  whereiis  st ininliit ion  of  the  houiolateral  peroneiil 
nerve  prothiees  an  extension  (inhil)itory  for  flexors,  oxcitntory  for  ex- 
tensorsV  Hy  takiiiif  ailvantaue  of  these  faets  further  proof  may  he 
supiilieil  that  inhihition  and  <M>ntraetinii  (.c.-iir  siiiinltanenusly,  as  shown 
in  l"i..'.  •-'14 

It  is  iiiipossihle  to  deiiionstiate  any  traee  of  inhihition  of  the  skeletal 


,^>M^. 


;'i 


'  I 


■;                ' 

'       , 

m 

I'lK,  Jl^.  J^litrntiKtMirs  ili.isiiam  illii>lraliiiii  the  nuchaniMii  hi'  rAipruial  iiiiu-i  \  iM'tn.  1  lu 
alTiTriit  lil.ii,  ^S)  from  the  skin  of  the  kg  ami  (S')  frcitii  the  Hex.ir  muscles  of  t|u  kme  un 
ham-lniig  nerve)  pass  to  ifie  siiinal  coni,  where  each  Rives  oil  a  hramh  which  diviues  into  two 
others,  of  whicll  one  in  each  case  S"es  to  a  motor  neuron  of  the  cMcnsor  nuiscif  s  I  /;  I  anil  the 
other  t.)  a  niot.,r  neur..n  ll)  of  the  Av\»r  muscles  (/-I.  Hrar.clR,-.  al-o  pass  aiross  llie  nn.lian 
line  to  similar  nioi.,r  reiirons  on  the  o|.|,.,sitc  side  of  the  c.nl.  As  in.licate.l  by  the  plus  an. I 
miir.is  siitn-,  ihe  allerent  stimuli  either  stimulate  or  inhilot  the  activilirs  of  tlu-  motor  neuron-, 
ihi    .i.urinioati.oi   ..f   llie   e\a.  t   ,  ti'ect   l.e-n  ;   a  fuiulion   of   the   ~v:ial-is.      I  I'rom    Sh.  rnouton.) 

muscles  hy  stimulation  of  their  motor  iiei\es,  thus  indicating  that  in- 
hibition is  dependent  upon  the  nerve  center.  Kurlherinorc.  since  inhihition 
occurs  aloiifi  wiili  llexion  of  the  antaL'oiiistic  muscle,  \vc  must  assiiiiM- 
that  the  afferent  impulse  on  entorinjr  the  spinal  cord  divides  into 
two  branches,  one  goiii!;  to  one  motor  neuron  so  as  to  excite  it,  the  other 
to  another  neuron  sn  as  to  inhiliit  the  toiiie  stiiiiuli  which  it  is  coii- 
-tantly  sc-ndinir  t"  the  tiiuscles  !  Fiir.  21'!  i. 

Since  the  seat  of  the  inhibition  is  in  the  nerve  center,  it  is  to  be  ex- 
pected that  impulses  transmitted  from  other  parts  of  the  nervous  system 


lik.i  ll'llni   \l,    INM  l(\  V'Hi'\ 


Hill 


than  th-  parlirular  l.-vl  oi'  that   rrlh-x.  uill  also  1..  able  to  nuliu-  th. 
inhil.ition.     In  th-  ••as.,  of  th.-  .l.-.Tol.rato  -at  tlm  cnti  l.o  "l-'..0"strat... 
bv  stimulation  of  tho  latnal  .•ohn.u.s  of  th«  spinal  cord;  nih.b.t.on  of 
the  oxtonsor  ,nus..l..s  ..f  tho  .•ll.ow  .ioint  opcnrs.  wl.i.-h  is  nil  the  more 
innrktHl  h.-.-ausc  in  s.fh  a  i.trpaintinM  thcs,-  n..is.-l,.s  a.T  n.  a  stat.  o 
hvpoHoni-itv      \Vr  shall  s...  lafr  aUo  that  thl•on^O.  tl.o  pyia.nulal  trn.-t 
impulses  tnav  ,lcs..,>n,l  f.o„.  tin-  .M.rl.rnni  uhirh  rx.T.Ms..  a  .luu-k.-d  in- 
hihitorv  influx......  ov.T  thr  .rtl.x  a.-tiviti.s  of  thv  .-onl.     Snn.la.ly  the 

inhil.ition  its..lf  iMMv  I,..  trnuiMMtr,!  by  i.n|.Mls..s  fion,  oth-r  soum's.  and 
the  motor  n-nron  tlius  thrown  fion.  a  state  of  inhibit io,.  into  one  of  eA- 
citation.  Tbis  fact  can  pcrbaps  best  be  ,lc.nonstfatc<l  by  cxeit.nK  the 
,.,.„t,al  .'ncl  of  the  hoa.olaleial  peroneal  nerv  (ubieh  pro,luces  a  rellex 
..xtension  of  tlie  le«  i  ^^hile  the  b-  is  bein-  hel.l  in  a  ib'xe.l  position  b> 
stimi.latioM  of  the  contralateral  i.enmeal  nerve.  This  v  ill  be  clear  from  a 
stn.lv  of  Ki-.'.  'JU. 

Snch  alternating  ex.-itatio,,  an.l  inhibition  of  an  active  motor  neuron 
serve  to  n,ake  it  possible  for  rhvtlnnie  .lis.harL-s  1o  occur  thrond,  jbe 
uetiron.  as  in  the  action  of  the  muscles  of  the  Ic'  in  walkmR  or  clt.r.njr 
the  scrntehinp  movetncnt.  In  or.b.-  to  insure  that  the  same  final  eom- 
,non  path  niav  be  ocenpicl  at  on.,  time  by  but  one  kin.l  of  stimulus  either 
inhibitory  or  excitatory,  it   is  further  of  iinportam-e  that  the  after-dis- 

,hai ("see   pa-e   SIO      of   the   first    stimulus   sboubl    be   .•apable   ot    l.ninc- 

rlinte^nhibition;  otherwise,  while  one  reflex  was  in  progress,  ,t  would  be 
impossible  t..  start  another  of  a  .lifT.Mcnt  type  employinit  the  s,.me  motor 
neuron  withmit  confusion  ..f  movement.  That  this  oc.-urs  eon  be  demon- 
strated  in  the  case  of  the  affr-diseharirc  of  the  flexion  reflex  by  stimula- 
tion of  the  proper  afferent  nerve. 

Tn  view  of  all  these  facts  it  is  probnble  that  the  seat  of  the  reeiproeal 
innervation  is  at  or  about  the  synapsis.  In  other  words,  the  synapsis  at 
the  termination  of  one  collateral  will  allow  a  siimulatins  impulse  to  pass 
to  the  cells  of  one  motor  neuron.  uhere;,s  that  at  the  end  of  another  col- 
lateral of  the  same  afTerent  liber  will  allow  an  inhibitiusi  impulse  to  pass 
to  an  antaironisfic  motor  neuc.n,  these  conditions  bein-   however,  readil.v 

interchangeable  and  thus  makin-.'  even   r.-ipid   rhytbmi mtraction  and 

relaxation  a  possibility. 

The  Action  of  Strychnine  and  Tetanus  Toxin  on  Reciprocal  Inhibition 
TTnder  certain  conditions  r,eiprn,-al  action  may  fail  to  occur,  as,  for 
ex.tmple   at  certain  staues  of  .tnirlnnn,  ,,o!so,iin,i  and  durin-  the  action 
of  tctanu.  toxin.     In  order  to  demonstrate  this  failure  of  reciprocal  ac- 
tion, it  is  necessary  to  examine  muscles  whi.-h  act  on  one  joint  only,  and 


.'   ' 

4 

.               i 

? 

\i 

m    I 

''k-     '  1 

ct           .     1 

J 


I       I 


Ax'' 
.■I    I 


S'JO 


Tin:    (I  NIIIM.    \1  IIVDI  S   SYSTi;M 


■f'; ' 

J    ill 

,1- 

;•■ 

r 

1 

it  i 


i^liiP:! 


I 

! 

1  1 

1!^ 

I'l 

i 

to  olisiTVc  tlii'if  Ixliiivior  wlioii  nil  n(Trri<nf  nerve  is  stimulated  wliieli  un- 
der (inliii.iry  eoiKlitidiis  ^v..lll<l  tlirow  tlieiii  into  iuliiliilion.  Sueli  a 
Iin|i.iiMtinii  Clin  iw  <ii>t;iin<'<l  in  the  liiiiil  liiiih  of  n  ih>\t  1>.V  euttintx  all  tlic 
nius.-lcs  flt.it  act  on  llie  i<nee  joint  eseept  tlio  vastus  erurciis,  which  in  a 
normal  aniniiil  invaiiahly  innjertroes  inliiliition  wIh'u  the  eentral  eutl  of 
the  intcniiil  saplienons  nerve  is  st  nnihiteil.  If  a  suitahle  dose  of  strych- 
nine is  injivte.l.  it  will  he  I'ound  that  stinnilation  of  the  interinil  saplieiioiis 

nerve,  in   i)laee  of  itiliihilion,  causes  iliaction   of  the  vastus  orureus 

muscle.     The  same  re. nit  is  olitained  liy  injection  of  ti'tanns  toxin. 

The  failure  of  the  reflex  inliihilioii  explains  the  symi)toms  produeed 
liy  these  sulistauces  It  explains,  for  example,  the  welMiUown  riiiidly 
extended  condition  of  the  lindis  in  stryr'hniiio  poisonintr,  and  the  dis- 
trossin;,'  symiitom  of  loehjaw  in  tetanus  infeetion.  In  this  latter  eon- 
dition  the  suffirer  is  suhjected  to  extreme  torture  with  every  endeavor 
that  he  maUes  to  open  tlie  jaw  for  the  purpose  of  taUiinr  food  or  drinl< 
Firnier  closure  is  the  result  lieeause  the  normal  inhihition  of  the  temporal 
aii<l  niasseter  muscles  does  not  occur.  Init  instead  they  heeome  exoited 
and  the  jaw  all  the  nn)re  fiiiidy  closed.  \ot  oidy  does  the  inhihition  fail 
to  occur,  hut  the  aliove  muscles  are  usually  in  i\  state  of  eonstant  hy- 
perexcitahilit.v.  which  it  is  inipossihle  for  the  patient  to  restrain;  indeed, 
whenever  he  attempts  to  do  o  the  opjiosite  oeeuis  anil  the  excitation 
lieconns  heiirhtened.  f'hloroform  acts  on  reciprocal  innervation  in  an 
opposite  way  froiTi  strychnine  and  tetanus:  iiairu'ly,  it  paralyzes  the  ox- 
citation  of  the  eontractin<i  muscles. 

Finall.v.  it  must  he  i>oiiited  out  tliat  this  im'chanism  of  reciiu'ocal  in 
•lervation  is  !>.»•  no  means  eonfmed  to  the  voluntary  muscles.  We  have 
already  seen  ih.it  it  occurs  in  the  case  of  the  myenteric  reflex.  Tt  is  also 
a  most  imiiortant  function  in  the  innervation  of  the  Mood  vessels,  dilata- 
tion in  one  vascular  area  heini:  accompanied  hy  constriction  in  another. 
These  facts  have  heen  already  sufficiently  dwelt  upon  elsewhere  (papre 

24:',i.     Sniiieti s  also  we  may  ha\e  ri'ci])rocal  action  lietween  differently 

netinsr  nervous  mechanisms,  as  for  example  m  the  ease  of  the  submaxil- 
lary irlands.  which  icspond  to  stinnilalion  of  the  clnu'da  tympani  nerve 
hy  dilatation  of  the  Mood  vessels,  an  inhihition  of  their  tone  oceurrins 
aloii!!  with  stinnilation  of  the  activity  of  the  gland  cells. 


CIIAI'TKU  XCIl 

INTi:i{.\» Tl(»\  A.MoN'd  UKFLKXKS 

A  Hiuah'  iv(l.\  a.'tinir  in<lr|Mn.l.Mitly  ..f  tin-  vr-\  <>f  tl.o  (•.•MtrM  ii.tvous 
syst.'iii  (liMN  not   iviilly  occur.     An  iifTcrcnf   impulse  on  cntciinir  llic  conl 
sptTii.ls  M.  iis  1o  involve  ii  liip.'c  Viiiictv  of  motor  neurons,  cmcIi  of  wliicli 
may.  Iiowever,  ]w  eNcil.'.l  tlirou-li  otlier  iilTer.iit    (il"Ts  jirrivinir  citJior 
frnni  other  re.'e|.tors  or  from  liisiher  nerve  i-enters.     Tlie  motor  neuron 
itself  nmy  tlierefore  l.e  ii   ptitliwiiy  oceupie.l  at   .lifferent   times  l.v  very 
.lilTerent   tyi>es  of  nerve   iniimlse.      llen.-e  it    is  niipropriiitely  ejillrd   Ihr 
fn>nl  rnmmnu   pnlh.  i.imI   its  netivity  iit    iin\    moment   must   depenil   on    tlie 
niituro  ofllio  viirious  .•iffen'Mt  impulses  tlial  lire  trnnMiiilt.  <1  to  it  tlirouuli 
tlie  synr\)>ses.     In  oilier  wonls,  an  eiiterinu'  afferent  filier  must  communi- 
cate in  tlio  cord  uitli  interntincial  patlis  vliicli  are  avaihilile  in  various 
(lejrrees  to  otlier  alTerent  fillers.     Since  it  is  tlirouuli  internnocial   paths 
tliat  tlic  impulse  is  tran-mitte.l  to  tlio  final  common  patli,  it  is  obvious 
that,  if  afferent   im)uilses  in   several   of  tliese   patlis  wore  eompetin-r  r 
tlie  same  time  for  tlic  possession  of  the  final  crmnion   path,  confusion  of 
movement  uoilhl   result  unless  some  provision  >v.ic  nia.le  whereliy  only 
one  kin.l  of  slimuliis  coiihl  l.e  transmitt.'.l  at   one  time.     "One  Iviinl  of 
stimulus  must  he  inhil.ite.l  ami  the  other  facilitated  in  its  occupancy  of 
tlie  final  common  patli." 

To  nn.lerstan.l  the  nature  of  lliis  int. ••--ration   of  the  ciilral   nervous 

svstem,  it  is  therefore  iiecissary  for  us  to  i si.ler  ihe  factors  which  <le- 

termino  u-hich  of  firn  mmpffliifl  iilTrrnit  !mpiiJ.'<r.«  xhall  ohttvv  pofmrxxwn 

of  the  filial  roiiniion  imth.     Let  us  take  th impetitioii  l.ctwccn  the  flexion 

reflex  and  the  scratch  r.tlex  of  till'  spinal  do;,'.  If  we  elicit  the  scratch 
reflex  and.  while  it  is  in  proL'nss.  apply  some  iineunns  stimulus  tn  the 
skin  of  the  hind  leu'  and  thus  indiu'c  the  (lesion  rellex,  it  will  ho  found 
that  the  scratching  movement  suhsides  and  the  flexion  movement  comes 
on  wilhout  any  overlappiiiL'  or  c<inl'usioii.  If.  however,  the  stimulus 
responsible  for  the  scralehin'.'  movement  is  n  strone  one.  and  that  ap- 
plied to  the  skin  of  the  hind  le-  a  feeble  one,  then  the  displacement  may 
not  occur  (sec  Fiyr.  '21(l~i. 

Tn  eonsiderin!?  this  inteu'ration  of  ri^flexes.  as  it  is  called,  we  must  dis- 
tinpuish  hotween  those  that  are  allied  and  those  that  are  antasonistie, 
and  wo  must  further  distinguish  between   reflexes  that   arc  simultanc- 

8:21 


m 


\- 


iM 


4fi 


t 

t 

1 

'ii! 

I  ' 


I    ' 


11  II 


8'2'J 


TIU;   <  INTHAI.    NIKMM  H   SYMb  M 


h 


M 


m 


! 

1'^ 

V. 

\ 

1     ' 

1 

1 

'' 

i 

,     \ 

« 

1    1  ■ 

1 

1     ! 

1 

1 

u 

='l 

' 

ouRly  PompcthiB  for  the  wimo  tiiinl  onmnion  pnfh  niul  thow  whieh  occupy 
il  siit'ccsNivcly. 

INTEORATION  OF  ALUED  REFLSXES 

IVrlmpN  the  sinipli-st  i\|)<riiii«iit  fn  show  tliis  is  perfornnMl  by  iisinif 
the  Norntch  roflox.  The  skin  nri«n  from  which  this  icHt-x  enn  ho  clicitid 
is  MTV  uitlrsprciHl  iMT  Kiu'.  '.M"'.  thr  type  <>r  vf\],'\  pn.<lii.'c.l  front  iiii> 
u'ivcii  nvcn  lirinir  in  •.'•■ni'iiil  tin-  snnit',  iilthonirh  "the  locnl  sign"- -that 
id.  the  point  nt  whii-h  tlic  nniiual  scrntohos  will  viiiy  accoidinR  to  tht 
point  stininlnlc.l.  If  thtii  «<•  tiikr  point  -t  in  the  ivllcx  scratch  nrca 
nnd  nppl>  to  it  a  stiiunlns  uliicli  is  jnNt  iiia<lci|uiitc  In  produce  any  reflex 
at  nil,  and  then,  wliilf  this  slininlns  is  still  in  pro^.css,  apply  a  similar 
siibliminnl  stinnilus  to  joint   /.'  .i   littl.-  rem     ed   from   it,  the  two  suh- 


l-iif  K  IiMtiani  .':  .«ii.»;  the  rcfl.x  arc«  invnln-.l  In  tin  mauli  rrflcx  Ko  «'>;1  A3  rr|ir<-»<ii! 
Ihc  affrr.i>t  11.  ..I  -  .Mpn.r.i.l  with  h.iir«  nn  Ihf  •Viii  ..f  llir  laik  aM'l  fl.ink  The  .iffrrrnl  m 
Pul.f?   arc   lr.in.mi(i".|   l.v    l>;ui-    IiIxt'.    aii.l   on   rntiri..!;   Ilir   ,  „rr,-|..-n.hn(,   jfgmtnt.   n(    llij   iplnal 

cIn'vM  in  .h.  lateral  cuiu.nP.  t,.  t.-rmuiatf  .im.larly  :„wm.<  <h.  .,1U  ..f  lh.-nK..,„  neurons  that 
iniKTvatr  ihr  m.is.lf.  of  th.  hi.l.l  lini'>.  Sinie  :ilf.r,.it  im, .„!>.■•.  ...■hmi!  (f™  'l«<-«''"'-  P"' 
licillarly  fr..m  iIh  .i..ii  .f  Hi-  La  (W  an.l  /.).  aK..  t.  riinr.alr  i.n  Hum-  n<-ur(in«  an.l  may  rxcitt 
Ihini  1.1  a  ilillcriiit  lv|it  i)f  a.  Ii"n,  tin-  nrntor  n<-\ir..M  ii  called  the  linil  comninn  lalh  (/-.C). 
(From   Sherringt..n.) 

liminal  stimidi  will  hceoino  cffii-tivf  iind  produce  n  typical  scratchinjr 
movement.  Tn  other  wcrds,  the  subliminal  stimulus  of  point  A  be- 
comes added  on  the  llnal  i-ommon  p.ntli  with  the  subliminal  stimulus  of 
point  R;  the  one  has  reinforced  the  other  nnd  produced,  therefore,  a 
simultaneovx  inieqratinn  of  nlUrd  rrflcrfn. 

The  receptors  from  which  these  mutually  reinforcintr  impulses  are  re- 
ceived need  not.  as  in  the  nbove  csan-plf,  be  of  the  same  kind,  similar 
results  lieiufr  olitaincd  by  stimulntion  of  receptors  of  widely  different 
Kiniis,  suc-li  ;i>  I'xtiTocc'ptois  nnd  pro|ii'ioccptors  t'sci'  pane  TS8).  For  ex 
ample,  if  a  stimulus  inadeiiuiili'  to  elicit  a  flexion  reflex  is  npplied  to  the 
iskin  of  the  lejr,  nnd  nnothfr  '^tiniulu';.  i««p]f  also  inadequate,  is  ap- 
plied to  the  central  end  of  some  deep  afferent  nerve  in  the  same  leg, 
then   the  two  subliminal   stimuli   will   become   effective   in   prodticinp   a 


INTKHAI  Tll)\    AMONW    BKriKXl.K 


«2:i 


flexion  in..vmH.t.     NcverthoU'ttn.  th.«  inoro  clonely  alli.d  the  rcccptorH 
arc  to  one  nnothcr.  the  more  cnsily  •I.k'h  nuiiimaticn  "ecur. 

The  mutual  r.iuf..rr.-m»'nt  uf  allif.l  r.thx.H  laMfn  f..r  n  short  time  aftt-r 
the  Htiinulation  ha«  bom  muov.d.  the  ph.t.«mo...,i.  h.i..«  now  known  as 
»uccf»m-e  integration  of  allied  rrfhs.>.  It  .-an  h,-  illu^trat.-l  also  m  iho 
east"  of  the  Koratch  nllox.  If  point  .1  on  the  nkin  area  is  .xeitid  with  n 
stimulus  that  in  itself  woul.l  he  inade<iuatc,  immediately  after  an  effet- 
live  stimulus  has  hern  .liM-ontiiiiie.!  at  point  11.  thru  thr  s.Talrh  mov 
merit  will  he  kept  up  sni..olhlv  although  it  will  of  eouihc  become  modi- 
li,.,l  ill  I.hmI  si«ii.  For  the  same  reason,  a  moviii(f  stimulun  applied 
to  the  seraleh  area  is  far  more  effeetive  than  a  statioiim y  stimulus  ap- 
plied over  the  same  eMeiit  of  area  In  sueh  a  ease  the  stimulus  that 
.xeilos  a  reflex  tends  by  its  oeeiipaiuy  of  the  nervous  pathway  to  facdi- 


^yi 


I  r«.    :\7 


SherritiRtoii.l 


tate  the  spread  alonjr  the  sam.-  patlnvay  »{  sueeeedin«  allied  stimuli; 
towards  such  it  lowers  the  threshold  of  -xeitability  ..f  the  reflex  arc. 
This  phenomenon  is  also  often  enlled  immrdialr  induction,  and  it  is 
by  no  means  confined  to  the  spinal  cord.  It  is  well  iUustri.ted,  for  ex- 
ample, in  the  ease  of  vision.     If  a  thill  line  drawn  on  a  white  card  be 

lu„|;,.,l  :it  so  th;it   it   TmIIs  mi  tli.'  >•.!-.•  «<  the  I i-tiv.^  tiehl  of  the  retina. 

it  will  not  he  seen  so  well  as  :i  dot  of  similar  width  which  is  moved 
Mirouph  the  sjiine  distaiiee  as  the  line. 

From  these  faets  we  see.  then-fore.  that,  when  two  allied  impulses  are 
l.,.i„i.'  tri.h«initte,l  to  tlie  final  .•..miMoii  pi.th.  tlie  one  is  likely  to  reinforce 
the  other.  ;.nd  thnt  this  tenden.-.v  to  reinforce  the  allied  impulse  is  main- 
tained for  a  brief  period  of  time  nfter  the  imptilse  has  been  removed.  We 
,,iay  novv-  proceed  to  consider  th.-  factors  which  will  become  operative 
in  dcterminintr  to  which  of  two  compctiiii.'  or  antngonistic  reflexes  the 
final  conmioi!  ))ath  "ill  hreome  iivailable. 


I  I 


u 


^■1 


U     1 


■    ! 


«i» 


\m 


S2t 


Tin:   ClA-IHAr.    NKHVorS   SVSTI.M 


4    - 


it  :{ 


Integration  of  Antagonistic  Reflexes.-  Altlioi^h  tho  iiliciK.inpnon  of 
imiiinlialc  in.luctioii  encouiii-;fS  iiitcffintion  of  allica  rcllcxcs,  .vi't  it  is 
fmiunitlv  siicc(-r(lf<l  l.y  one  of  siirif^sivr  induction,  in"  wliioh  just  Ili.> 
,.,.posito  ("oiidilions  ..ccnr;  the  iTsistanco  in  tlic  rofl.'x  pjitlnvny  bpcomes 
lowered  for  a  type  of  moviniont  antapronistic  to  that  Avliidi  tirst  occu- 
picil  tlio  roflox."  To  iindi-pstand  clearly  what  relationship  this  hears  to 
innne.liate  iii(liH'ti..n,  it  may  he  well  to  take  the  instanees  in  which  those 
phenomena  ni.ply  in  the  case  of  vision.  Wlwu  the  eye,  after  .larkness, 
is  sml.lenlv  iVirected  to  a  liu'ht  an.l  then  closed,  there  remains  a  l.ritrht 
ima-e  (positive  after-effect^  of  the  li-ht :  hut  if  the  li«ht  is  looked  nt 
for  some  time,  then  on  closinir  the  eyes  it  will  he  seen  as  a  dark  pat- 
tern fneirative  after-effectV  In  the  former  instance  wo  have  an  exam- 
ple of  imniediate  induction,  in  the  latter,  one  of  snccessive  induction. 

Tn  the  spinal  aiuiiial  snccessiv.'  induction  is  demonstrated  with  equal 
ease  hy  usin-r  two  rcllcxcs  th:'  are  of  a  more  oi-  less  antauronistic  charac- 
ter fi.r  eNaiiiple,  the  llcxion  redox  ;iiid  the  knee  jerk,  or  hettcr  still 
the  crosr  .d  extension  rcllcx  and  the  ih'xion  reflex.  If  we  elicit  the  k,i(>c- 
jerk  in  a  spinal  do;:  at  regular  intcivals,  with  stimuli  of  ei|ual  intensity, 
the  extension  movements  flhc>  ki.-ks~)  will  \w  ai)i.roxim.itely  equal.  If 
now  we  apply  a  nocuous  stimulus  to  the  skin  of  the  foot  and  so  throw 
the  loc  into  fh'xion,  il  will  he  found,  alter  the  flexion  movement  has  dis- 
appeared, that  the  knee-jerk  is  much  more  itrononiu'cd  than  previously. 

Similarly,   if  we   elicit   tin-   cro^scd   extension    rellc\   l.y    i uons   stimuli 

of  equal  intensity  applied  to  the  opposite  limli.  the  ,.stcnsion  Tnovcnn-nts 
will  1)0  approximately  eipial.  Hy  now  throwins  the  liml.  cNliihitinir  them 
into  the  flexion  icilex.  the  cxlcnsco'  tiio\«  incnts  will  of  course  disappear, 
lint  aft.'r  the  llexioii  has  hecn  .lisconliiiucd.  they  will  rcajipcar  with 
marked  intensity. 

These  facts  show  \is.  then,  that  after  the  linal  common  I>a1h  luis  hecn 
ooeui)ied  hy  a  reflex  of  oiu'  lyjic.  it  hoconics  more  availahle  to  n  reflex 
of  an  oi.i>osito  type.  In  oilier  words,  it  is  evident  that  if  the  two  op- 
posite reflexes  are  constantly  o(.m|)etini;  with  each  other  for  possession 
of  the  final  oommon  i)ath.  they  will  tend  alterniitcly  to  occiijjy  it.  thus 
hriuKinir  ahoiit  a  rhythmic  movement.  Such  is  the  mechanism  involved 
in  walkin--':  the  loir  is  lifted  from  the  -rround  (llcxi'in  i-cflcx")  :  it  is  then 
hrou<rht  on  the  -rroun.!,  and  the  me<'hani.-al  itush  '.nvcii  to  the  iilantar 
sTirfaeo  of  the   foot  hrin<:s  out   the  extensor  thrnsl.   the  appearance  of 

which  is  -rroatly  fa.  ;!italcd  hy  the  fact  that  ii cdiatcly  hefore  the  llexion 

reflex  oi'eui)icd  the  fiiuil  common  jiath. 

Other  Factors  Which  Determine  the  Occupancy  of  the  Final  Common 
Path.'  Hesiu.-s  immediate  and  suc,m-,s1\  .■  inductiori,  several  other  far- 
tors  nffoet  tiK  relative  availahilily  of  the  reflexes  to  afferent  stimulation. 


iNTi.KAiTioN-  AM<iN(i  ui:Ki.i;.\i;s 


Iiii])()rtiiiit  aim 


,-  thcs,-  is  fatigue  of  the  nllrr  an-  for  <i  porlh 


iif  stimulus.    Many( 


■liavai'tcristii's  dilTcri'ii 


•iliir  hiiiil 
tiaU'  ri'llt'x  faliauo  from  fatiuMie 
„f  a  u.Tvo  as  oLscrvnl   in   an   isdat...!  luTVo-niMs  '.   ,,r..parati..n.     Tho 
most    important   of  those   ,lis1iM..r„isl,inff  f.atnrcs   arc   as    follows:      (1, 
Tl.o  fali-n.r  .'om.'s  nn   inl.'rniittontly :  tlius,   when   t'u'  il.'xion   r.llcs   is 
pnrsistontlv  olL-itcl,  tl,o  first  si-.m  of  fati^M.o  is  an  irro-.'ular  <Whno  in 
the  flexion  movement   f..ll,Ave.l  by  its  entire  .lisappearanee   for  a  short 
time      These  lapses  hee„me  move  and  more  fre.,nent,  nnt.l  at  last  com- 
plete  fatijrne  sets  in   an.l   no  flexion   oeenrs.      (2)    Hellex   fatiun.c  soon 
passes  ofT     (■^)  Tt  app-ars  earlier  for  ueak  than  for  strong  stnnnl..     (4^ 
The  n.ovement  pro.lu.v.l  hy  the  ,vtlex  a.'tion  may  also  ehanire  u.  eharaeter 
dnrin-  reflex  fath.'ne;  thus,  the  heat   of  the  serateh   rellex   may  l.eeome 
slower  an.l  less  stea-lv  an.l  the  font  he  less  aeeurately  dire.-te.l  to  the 
spot   stimulate,!.     The   l-.ens   of  the   fati.me    in   the   rellex   are   ean   not 
ho  the  motor  nenron  itself,  for,  af  -r  this  has  heen  eomplefly  fat.sue.l 
hv  stimnla.ion  of  the  serateh  a,v.     the  sanu-  museles  nmy  .pute  rea-l.  y 
he  thrown  into  a  perfeetly  nor.nal   fl.'xion   rellex  hy  sinnnlation  of  the 
skin  of  the  hind  le-.'. 

It  is  evident   that,  when  two  reil.'x<>s  are  eompetnin-  w.1h  eneh  other 
for  possession  of  the  same  linal  eommon  path,  the  one  that  l.eeomesla- 
thmed  will  he  mastered  by  the  other,  espe.-ially  sinee  at  the  same  t.mo 
s,n-eessive  indnetion  will  be  well  .leveloped.    Thus,  ordinarily  the  serateh 
veflex  is  mueh  less  readilv  .'lieited  than  the  llexion   rellex,  and  if  both 
ave  exeited  at  the  same  time  the  latter  will  i.revail:  but  if  the  llexn.n  re- 
flex  is  kept   i.p  nntil   it    slums  si-.nis  of   fati.-ne.  then   by   simultaneous 
exeitation   of   both    reflexes  the  serateh    reflex   will   obtain    the   mastery. 
\no1her  importani    faetor  is  thf  relative  strength  of  Ihr  ronu>,tn,u 
hnpuhrs-     This  .lepends  partly  on  the  nature  of  ih-  reflex  and  partly  on  the 
intensit  V  of  the  stimulus.     He^'anl !.,•_'  the  ,u,tun  uf  >h,  n  th  r.  it  is  ,mporta.,t 
J,  ,,.„,„„,,„.,•  ,hat  erosse,!  reflexes  are  usually  less  easily  obtained  than  homo- 
hiferal   ones    but   of  still   ureater  import anee  is  the  spee.es  of  reflex- 
,I,at   is    whether  flexion,   serateh.  extension,   ete.      The   reflrx    movements 
produeed  hv  noeuons  stiunili   f noeie.ptive  refl.'xes^,   always  take  pnMM.denee 
„f   those   produeed   bv    oth.M-   kinds   of   stimuli:    or.    to    put    it    n,    ..ther 
words,  "noeieeptive    reflexes   are   prepotent    in    their   oe.Mipaney    of   the 
flnal  e.muuon  iiath"     fSherrinuton'- '. 

The  best    known   example  of   a    ,u,rir,  pllrr    r,  ,h  r   is   the    flexiou    reflex. 
Tts  movement  is  o.e  prodneed  with  the  intention  of  nn.ovincr  the  ..t,mu- 

lated  portion  of  the  hodv  fnan  the  son, of  th.-  stimulus,  all  stimuli  whieh 

produee  it  b..in._'  sueh  as  would  elieit  pain  in  an  intaet  animal,  or  ,f  por- 
<,i,to,1  i„  e,.use  some  dama-re  to  th,.  skin.  Tn  eontrast  to  sn,-h  noei.-eptivo 
reflexes  we  mav  take  those  xvhi.di  are  eoneer.u'd  in  mamtanun..'  the  e.'n- 


i'i 


-  iM 


3   i;  ■  i' 


826 


Tin:  CKNTKAI.  NKKVOrs  SYSTf.M 


m 


tor  of  gravity  of  tlie  hody-postural  reflexes,  as  they  are  called.  The 
l.pst  type  of  this  retlrx  is  tlic  kiioe-jiTk,  another  Rood  exiimple  boiiif?  the 
.xteiisor  thrust.     The  .serateh  reflex  contains  a  certain  element  of  the 

ii.M-i.M'ptivc  ill  it.  ■•m.l  of  the  siiniilcr  reflexes  it  eonies  se<'on(l  in  its  clfiiin 
on  the  final  common  path.  In  brief,  then,  in  reflexes  wliieh  in  an  intact 
animal  would  cause  tin-  sensation  of  pain  and  prohaMy  some  reflex  ae- 
tivilv  of  the  voeiil  oi-ans.  we  jiel  in  llie  spinal  animal  a  reflex  flexic.n 
moveiiirnl  of  the  |)ait  stimulate.l  with  ihe  evident  object  of  removinu- 
that  part  from  the  stimnlatiiifi  agency.  This  reflex  flexion  secures  pos- 
sessi,,ii  of  the  final  eor.imou  jiath  whatever  other  reflex  may  at  the  time 
bo  occnpyin-r  if.  Thus,  if  the  animal  is  serntchinir  itself  and  somcthiiiL' 
oeenrs  to  hurt  its  foot,  then  immediiitely  the  seratchiiitr  movement  will 
<.'i\  ('  iilacc  to  one  of  flexion,  and  so  on. 

Some  iiifetrr.itioii  between  distant  reflex  ;u-es  in  the  nervous  system 
is  to  .'i  eerfaiii  rxleiit  nil  aiiplieatiuii  (  '  the  prinei])!.'  of  reciprocal  in- 
hibition i.f  the  musc'les  iiiovinu'  a  joint.  In  this  broader  intepration  the 
inhibition  affects  mor.'  removed  fields  of  leth^x  aetivify  so  as  to  harmonize 
the  activities  of  one  part  of  the  animal  with  those  of  every  other  part. 
The  manner  in  wliidi  the  stimulation  may  spread  alonir  the  various 
available  i>ath\vavs  als.i  i]e|)eii(ls  on  lh<  slrniiith  of  Ihe  nll'o-Dit  hii 
piilsis:  If  a  \.'iy  feeble  stimulus  is  ajiplied  to  the  skin  of  the  leir  in  a 
spinal  animal,  the  reflex  will  lie  represented  only  by  a  sliudit  contraction 
of  the  inner  ends  of  the  hanistrinsr  muscles  .\s  the  stimulus  is  increased 
in  strenfrth  the  reaction  will  spread,  nntil  at  last  it  involves  all  the 
flexors  in  mnfm  Hon  and  the  anta^'onistie  extensors  in  iiiliihitwv.  Tf  it  i- 
still  further  itiereasod,  the  flexion  movement  will  be  aeeonipanied  by  an 
extension  of  the  muscles  of  the  op).osite  hind  limb-  the  crossed  exten- 
sion reflex.  Further  increase  of  the  stimulus  will  cause  the  reflex  move- 
ment to  s|,read  to  the  anteiioi-  extremities,  iii\olvin-r,  first  of  all.  the 
fore  limb  of  the  same  side  '  extension  at  the  elbow  and  contraction  at 
the  shoulder^  and  then  that  of  the  opposite  side  fflexion  at  the  elbow 
and  extension  at  the  u  list  .  .\  \eiy  powerful  stimulus  ajiplied  to  the 
hind  limb  will  even  si)r<.n<l  to  other  more  distant  muscular  ffroups,  such 
as  those  of  the  neck,  causing  a  turnin-r  of  the  head  to  the  side  stimu- 
lated, opening  the  mouth,  etc. 

This  siircad  or  irradiation  oi'  tlie  r.'llex  in  tlie  spiiml  eord  c.-in  not  t.r 
entirely  rxi)lained  on  anatomie  trrounds.  and  must  dei>eiid.  therefore, 
upon  varyiiiu'  resistaii<-e  to  tlie  flow  of  the  afferent  impulse  to  dirferent 
motor  neurons,  some  of  which  it  exeites  while  otlii^rs  it  inhibits. 

T},(>  iipof-ssity  for  adjustable  resistance  to  the  transmission  of  different 
alTercnt  stimuli  on  to  the  final  eominon  iiatli  Ijeeomes  evident  wlieii  we 
ii'member  that,   not   only   arc  there  about   fixe   times   ;is   many   fibers  en 


INTERACTION    AMONO    KEKI-EXKS 


827 


tering  the  cord  as  motor  fibers  leaving  it,  but  also  that  each  afferent 
fiber,  after  its  entry  to  the  cord,  gives  off  several  collaterals,  each  of 
whicli  runs  In  some  nerve  eentei-  in  the  eonl  (see  Fl^'.  20<). 

Certain  conditions  mny  l.real<  down  the  path  alona;  which  the  impulse 
passes;  for  example,  at  a  certain  stage  in  the  action  of  strychnine  al 
pathways  beeon.c  ..pened  n,..  su  that  the  reHexes  whieh  ordn.anly  do  not 
!„.cur  together,  a.-t  sinn.ltane.msly.  with  the  n-sult  that  a  typical  convul- 
sive movement  is  produced..  Strychnine,  as  wc  have  already  seen,  also 
interferes  with  the  sorting  out  of  the  impulses  into  inhibitory  and  ex- 
citatory, so  that  no  reciprocal  action  occurs. 


'*f 


Js» 


I  ;:    ^1 


11 


iiM    ' 


:!       11     i 


J, 


1      i 


S  1      i 


('ii.\rTi;i;  xciii 

TllK  TI:NI)<»\  .IKIJKS.  SKNSOI.'Y   TATIiWAYS   IX 
Hl'INAL  (OKI). 

Ccrtjiiii    |■(■^|ll.|ls.•s  iiri'  cit'  imiiort.Mii-c  biiirt'ly   Ik'i-musc  of  tlicir  clinical 
a|i|)li.Mtioii.     or  .^iviil.'st   iiiliiCNl   in  tliis  conn.'ctidM  iirc  tlio  tendon  y-'A:-. 
Tlic  l<M-!iti"n  ol'  til.'  sense, i\   i.alliwavs  in  llic  s|>inal  mra  nlso  .Icnninds  at 
tcntion. 

The  Tenflon  Jerks.     Onc  dt"  t!ic  ni^st  iiiii>nitant  icII.-ncs  for  .liairnostic 
liiirpo-^'"*  i-^  'liat  known  as  llic  I,  ,i< , -j,  rk,  wliicli   is  clicifcl   in  inan  by  »\> 
plyinir  a  sniavl  lap  to  tlic  patdbir  tendon  of  a  person  who  is  sittins  on 
n   Iiiuli   sto.>1   or  taMe  so   that    llie   .ioint    is  i.assivelv   flexed    and   llie  ]<■■_' 
Imn-s  loosely  from  llie  kni-e  joint.     In  this  i)osition  tlie  extensor  nitiseles 
nre  pnt   sliuditly  on  Ike  streteli.  and  "ken  tke  patellar  tendon  is  striiek. 
tkese  iiinseles  eontia.'t  and  .'anse  tke  leir  to  ke<-oMie  .'xlcnded  as  in  kiek"- 
in<r.      Tkis   relkx.   as  ue   kave   s,.cn.    is   also    rendily    elicited    in    spinal 
animals.      Its   importance    from    n    clinical    standpoint    depends    on    iho 
f;ict  tliat  if  may  lie  altered  not  only  in  vari.-ns  irenernl  conditions  of  tke 
kody,  kut  also  wken  any  i)atkolo-,'ic  ci>ndition  disturks  tko  pnntintiity  of 
tke  relies  arc  com'eriied  in  nianitainiiiir  the  tonicity  of  tke  extensor  mus- 
cles of  Ike  tliiuk.     Tk.>  centers  involved  in  tkis  arc  arc  situated  nkont 
tke   tkird   or   fourlli    Iniiikar  s.-jmcit.   and   tke   afferent    impulses   come 
partly  from  tke  antaironistie  flexor  muscles  ami  partly  from  tke  extensor 
nuisele  itself.    Akolition  of  tke  rellexes  may  tkerefore  l>e  produced  eitker 
l.y  neuritis  inv.ilvini.'  tke  afferent    likeis  or  myelitis  affeetin;.'  tke  i:ra\ 
matter  of  tke  cord.     Tkat  certain  of  tke  atVerent   imiMllses  cmie  from  tke 
kamstriii!.'  muscles  is  skown  liy  tke  fact  tkat  wlien  Ike  central  end  of  tke 
cut   motor  nerve  of  tke  <'\teiisor  mnsi-les  is  stimulated   electrically,  tke 
knoe-jcrk  hecomes  muck  less  evident,  a  result  wkieli  is  also  oktained  ky 
massairinir  tke  muscles. 

Altkouirk  suck  facts  skow  dearly  tkat  tke  knee-jerk  is  of  reflex  na- 
ture, yet  tliere  ai'c  (lifficul'ies  in  exiilaininir  Ike  exact  meekanism  h\ 
wkick  tke  tap  to  Ike  tendon  proiliices  tke  mus.uilar  contraction.  Tke 
ekief  difiii-ulty  is  in  acciuhlin-  for  Ike  in-oinptness  with  wliich  the  i-oiitrae 
fioii  occurs.  Ike  kilent  i-'''''"'!  ''ein-r  very  m\ick  skortev  tk.nn  tkat  of  suck 
rellexes  as  tke  flexion  or  even  tke  .•onjuni'fival.  Tke  total  latent  period 
(if  tke  kiicr-jerk,  as  .iud^'ed  ky  tke  time  elapsing  l.etwocii  npplyinj,'  a  tap 

828 


TI;N1>i>\    .IIUKS;    SF.NSdUV    l-ATIIWAYS    IN"    SI'INAI,    (  Olil> 


S-J!) 


to  tlio  tendon  iinil  tli 


ctrir 


I'lSllDIISC    ( 


.lis.i-,<'(l  ill  tlio  vastus  iiitiTuns 


imisclo  l)v  the  stiiii>f  iraU  iinoiin'l"'!-,  was  ion 


11(1  liv  .Tollv  ill  tlie  spinal  cat 


to  be  0.0055  of  a  sccoi 


.1.  \vl 


icrcas  iiicasui 


vi\  in  tlie  same  way  the  latent 


period  of  tlie  ttexion  rellex  was  foiiiid  to  I.e  just  twiee  as  loiij: 


,  O.OlOti  ol' 


a  seeoiid.     Tliese  differeliees  were  v\\ 


lained  l.v  Jollv  as  indicating  that 


flexion  leflex   involves  three  a 


the  knee-jerk  is  a  sinii)le  reMex,  involving  hut  two  neurons,  whereas  the 

iiid  tJKrel'ore   has  twiee  as  lonsr  a  latent 

tlie  total  latent  period  the  time  oeeiipied 

CI    IS  and  the  time  lost  at 

re  a  fiirure  iriviie,,'  the  lime 


period.     \'>y  suhtraetiiii.'  I'limi  me  loiai   lan 
ill  the  traii>iiiission  of  llie  impulse  aioiis;-  th 


the  atVereiit  and  ell'erenl  iiervi 
lost  in  the  syiiiiiisps  lietwcen 
ealled,  was  found  hy  .lolly 


i'iidiii'_'s.  we  seen 


11 


le  neurons. 


Th 


iiiajise  time,  as 


it  is 


to  he  (l.(l(»-21   of  a 


.f  a 


for  the 


AKiii   reilex. 


lid  (t()(M:i 

milar  results  in  man  hy  the  same  method. 
Some  authors,  parlieiilaily  (iowcrs,  do 


seeoiid   for  the  knee-jerk 
Siivder  olitaiiie<l  somewhat 


not, 


liow  ever 


I'l-k  is  of  the  liallin 


to    a    colli  I  aet  ion    o 


meeh 


aiiieal  slimiihitiiiu 


helii've  that  the 
f  ,1  simple  relle\,  nut  .■xphiiii  it  as  heinsT  due 

ahoiit    h\     direct 
■xeitahle  eoiiditinn 


f    the    cxlen  nr    muscles 


1. 


uht 


ilr  the  iiiii-clc  is  in  a  hy|)'.M( 


as  a  resii 
piittinsr  tl 


It 


r  a  rellex  iiicrca 

■xtclisor    lllllsi'leN 


,e  In   its  tonicity.     (1 
on    the   stretch    am 


ieliev( 


■s  that  hy 


1    the   haliistrintr    miischs 


tlio  relaxed  eonilition.  a 


iff. 


i1    im)>ulses  are  transini 


tied  to  the  eord 


uhieh  oxeite  tlie  elTereiit  neurons  of  the  ex 
them  into  a  hyiiertoiiic  condition,  dnriiii;  w 


tensor  muscles,  so  as  to  thro  v 


doll  directly  excites  a  i-o 
eoiint   once  and  for  all    tor 


liicli  the  tai)i>iliir  <'f  the  ten- 
traction,     (tf  course  this  h>tiothesis  would  ac- 


tl 


le   remaika 


hlv   short    lat( 


■k    lint  on   the  other  hand   it    haves  iis  i 


iiain'   I 


of  the  knee- 

ilr 


lifliculties  to   exlilain; 


eh',  for  example,  as  the  fact  that,  although  tapi.in-  the  tendon  produces 

the  muscle  itself  has  no  eft'eet. 


the  .jerk,  similar  tappiiiir  ol 

The  elVeetive  stinnilns  of  t 
tension  to  which  th 
latioii  of  I'eceiitors  ii 


le   .|« 


rk 


slight  11 


increase  o 


'xtensor  luu 


itself  is 


suh.ici'ted,  ■'111 


f  the 
d  not  a  stimu- 


Ihe  tendon,  for  it  still  oeeiirs  after  the  tendon  has 
hcen  denervated.     The  importance  of  ihe  relationship  of  the  hamstrinsi 
nerve  to  the  knee-i 
action  ;  thus,  w 


rk    1 


iceomes   evii 


lent 


in    eonnei 


tioii   with   reciprocal 


hen  tlie  llexor  is  eoiilraete.l,  as  in  the  flexion  reflex,  the 


-.)erl 


disai 


tpears  i  ]f,\ 


ire  S14\  whereas  \\  hen 


en' 


it  is  auirmciitcd. 


Whatever  its  nature  may 


1h(>  knee-.ierk  is  o 


list  film'  nerves  are 


f  value  liccansc  of  the 


case  w 
ell 


ith  -which  it  can  ho  altered  not   only  hy  conditions  a 


ffect 


1112 


rellex   are   coiiceruei 
central   nervous  system, 
known  as  n  injurt't  i)i<  iit. 


1    lull    also   hy  ehninres  o<-curriii! 


The  hesl    known   of  these  conditions   is 


plsewl 
nditi 


lere   in 


the 

the 

that 


make  some  voluntary  muse 


This  is  hrouirlit  ahont   hy  liavinu'  the  paiieiit 
nlar  effort  at  the  ir.oment  that  the  tap  is  ap- 


it- 


I     il 


i 


4'^'  'I  Mi  f; 


^!llM  i  -Ml'! 


S:{()  TIIK    IKNTRAI.    NKKVOrS   SYSTtM 

plied  to  the  teml..!..  If  this  vluntaiy  effort  ooiucides  in  lime  with  the 
lapping'  of  th..  tcn.lo,,.  th.-  ktioo-jork  will  lu-  found  much  aURmonted;  hut 
if  the  two  events  do  not  noourntoly  coincide,  we  mny  find  instead  that 
the  knec-iork  is  .liininishc.l ;  that  is  to  say,  we  may  have  positive  fol- 
lowed hv  nesiativc  ,viufn,r.-meiit.  The  most  usual  way  of  having  the 
patirnt  make  this  voluntary  etVoit  is  to  ask  him  to  lock  the  fmtjers  ol  his 
two  hands  together  and  then  at  a  triveii  signal  try  to  pull  the  locked 
arms  apart. 

Similar  reinf..iTement  may  also  he  produced  hy  the  application  of  a 
stronir  sensorv  stimulus  in  some  distant  part  of  the  nervous  system,  as. 
for  example,  hy  imllinir  the  hair  or  piii.'liinsj  the  ear.  Accurate  work 
,m  the  time  relationship  hetwecn  the  reinforeinn  act  and  the  tap  on  \\u- 
tendon  has  shown  that  the  knee-jerk  is  most  marked  when  the  tap  ac- 
.Miratelv  corresponds  with  the  voluntary  effort  ov  sensory  stimulation. 
It  then  .|uicklv  declines  and  an  inhihitory  inlhicnce  appears  in  ahoiit 
0.3  to  0.6  of  a  second,  immediately  after  which  it  hecoraes  pronounced 
asrain,  gradually  fadiiur  off  to  he  no  longer  evident  in  ahout  1.7  of  a 
s.MM.nd;  that  is.  no  .•lianire  from  the  normal  will  he  found  in  the  knee-jerk 
in  ahout  l.i")  of  a  second  after  the  reinforcing  act  (Lomhard'V 

Manv  explanations  have  heen  offered  of  the  mechanism  involved  in 
this  reinforcement.  The  most  commonly  accepted  is  that  it  is  due  to 
the  ov..rtlow  of  impulses  from  other  parts  of  the  nervous  system,  par- 
ticularlv  the  cerehrum.  upon  the  reflex  arc  concerned  in  the  knee-jerk. 
During  voluntar^  effort  the  cerehral  impulses  discharged  dowm  the  spinal 
cord  pass  not  onlv  to  the  neuron  for  which  they  arc  intended,  hut  ir- 
radiat..  or  spread  to  otli.-r.  .■vcn  far  distant,  neurons,  thus  adding  tli.'ir 
effect  to  that  of  the  afferent  impulse  entering  the  cord  locally.  The  suc- 
ceeding inhihiti.m  may  he  assumed  to  he  due  to  successive  induction  (see 
pajrc  s">4).  It  is  difti.'Ult  to  offer  din-ct  exiu'rimeiital  pr.n.f  in  support  ot 
the'^  explanation,  hut  indirect  evidence  is  furnished,  in  so  far  at  least  as 

the  augmentation  is  co rned.  Uy  the  results  of  the  experiments  which 

we  have  alreadv  descrihed  concerning  the  integration  of  allied  reflexes 
ipa-e  S22i.  To  these  ini-ht  he  added  the  well-known  fact  that  the  simul- 
taneous application  of  two  suhliminal  stimuli,  one  to  the  cerehral  cortex 
and  the  other  to  the  skin  of  the  cn-respondiuir  hody  area,  may  call  forth 
a  contraction  of  certain  groups  of  muscles. 


.'      1 


AFFERENT  SPINAL  PATHWAYS 

The  nalin;  of  tl,(  impulses  InuLs-mittcd  h,f  the  vnrhus  afferent  path- 
ways in  the  spinal  cord.  We  have  seen  that  the  sensory  impulses  travel- 
ing from  the  periphery  to  the  spinal  cord  group  themselves  into  three 


'  I 


•n  MM  I.N    .II.KKS:    SKNSOltV    I-ATIIW  ays    in    SI-INAI.    (OHI' 


<11 


olassrs:  protopnthip.  epicritio,  and  deep  or  niuscuhir.  It  is  important 
now  f..r  us  to  consider  what  l.ccomcs  ..f  each  of  those  impulses  after 
enterinir  the  spinal  ('..nl.  for  there  is  al.undant  evidenee  that  they  travel 
up  to  th..  l.raii.  hv  dilTeivMt  patlnvays.  This  evidenee  is  furnished  partly 
hv  ex.imination  of  th rd  nf  i.iitients  who  during'  life  exhibited  per- 
versions (.f  the  skin  sensations,  and  partly  l.y  prodneini;  exi.erimental 
lesions  alTec'tin^t  dilTerenl  j.i.rts  of  the  spinnl  eord  in  animals.  Tn  the 
disease  svrinsromvelia,  foi'  exami.le.  enlai-ement  of  the  eentral  .'anal 
of  the  spinal  eord  eanses  rui.ture  of  eertai.i  of  the  traets  and  a  eonse- 
(pien 


it  disintegiation  of  the  skin  <ensa 


;ind  temperature  disappear 
-cnsiition  remain. 


\vl 


tions:  that  is,  the  sensations  of  pain 
touch  and  deep  muscular 


th 


creas  those  o 


dr.  fii>iii  the  experimental  sidi'.  if  we  make  a  lateral 
hemiseetion   of  the  spinal   eord.  then   after  recovery,  so  far  as  wc  can 


studv  it   in  a  dumh  animal,  we  shall   He  a 


iilile  to  sliow  that  certain  sen- 


itjons   have   (lisii|ipe.'ii 


'd     wherciis   others   remaiii. 


II 


evident,  h 


ever,  that  we  must  .judsre  l.y  o1..ieetive  and  not  hy  sul.jeotivi  phenomena 

id   our  results  are  only   approximate  and  very 
Iniporlant  i trilnitioiis  to  this  subject  have 


01 


these   exiieriments, 


liable  to  misiiiterpretati 


llv  been   made,  particularly  by  llohnes^'  and   by   Collier.'  on  sol- 


diers wounded  in  the  spinal  cord. 


Suiiimiiit;-  up  the  res 


\ilts  olitaini'd  \<\    tiie  earliei'  iiivcstii:ators, 


n 


Scfpnird  some  sixty  >ears  ;iiro  s 
iide  produced  tin-   rollowiiii;  results 


.tated  that  he 


I'tion  of  the  cord  on  one 


(n 


paralysis 


of  voluntary  ntotion 
lie. 


<f  the  same  side:  i2~i   jiaral.vsis  ot'  vasomotor  control  on  the  same  sid< 

1  :  i:!''  anestliesin  for  all  kinds  of  sen- 

the  side  op|>ositc   to  that   of   the   lesion: 

lied  liyiteiesthesial  on 


that  the  limb  is  liottei'  than  iiornni 


satioii.  except   niuscular  ^eiiv 


1 

tlic 


diti 
si(h 


it'  heiu'hteni  1  skill  scnsilivit\    ^  ca 


as  till 


le 


-ion.  wi 


111  tlie  exception  of  a  narrow  strip  o 


f  ski 


Tl 


)irespol 
lese  lesu 


dinii'  to  the  ..esiiiienl  at  which  the  d  is  c\l 


1.  which  is  anes 


.theti( 


lis  indicate  that  in  ■ieiier 


\\  the  skill  sensations  of  pain,  touch. 


temper.-itllie   cross   oX'el' 


to    til 


other  siile  s 


hortlv 


after 


Ibeir  eiitrx' 


into    the    coi-d.    luit    that    the    deep    i 


iiiuseiilar  sensations   remain    in   larire 


part    uncro 

1)1     SU| 


il.      M 


ire   reet 


lit    ex|>eriiiieii1al    ,'1ihI    clinic:i 


il    iiivestiiratioii'- 


.Il 


)pol 


t   M 


row  ii-Se(|n;irii 


conclusions. 


IJansoin  li;ls  rcceiitl.v    slio 
iiitain    both    meduUateil    am 


tii:it   the  .■itTcrelil    loots  ol'  the  spinal  con 
d    iionmedullaleil    nerve   libers,   and   he   be 


kcs  that   the  former  transmit   the  epicrilie  sensations,  and  tlu 


the    jtrnlopa 


th 


Hv   Iracinu:   those   difl'cren 


latter 


kinds   of   libers    into   th 


spinal  cor<l,  he  found  that  the  iioii 

then  1 


ilullated  lie  in  I.issauer's  tract 


;>ass    ill 


I II  IK 


to  the  substantia  uelatinosa  Ko- 
ili,  which,  therefore.  ai>pears  to  be  the  nucleus  for  tiie  reception  of 


iiie  or  two  set"inents  ami 


tiie  ])rolopathic  impulses 


\\h 


111     i 


"i 
if 


If;  I 


^ 


i:^* 


8:Vi 


Tin;   ClATUAI.   MUVdl  S   SYSTIM 


ti'HlM 


V. 

11  ■ 

i 


1 
'  1 


S,„ tl,..  r.H,.x  ,M-tivili.s  «l.i.-l.  lM.-.m,.>  cv.'it.'.l  l.y  tli.'s.-  tuM-icoptiv.' 

i,„,,nls..s";,ro  IIm.s,.  .•lU.si,,.,'  n    risr  in   1.1 1   i.ivss,.n.     iwrssur  ,„.puls,.s. 

This   .•c.nclnti.m   brUvni.   I i.M.|.ti\r    iu.|.uls..s   .-in.l    Hh's,.   iillrHint,'   tl.o 

v.s.M.i.r  >vll.s,s  Ims  ,.r..,H!.t,.,l  Umms,,,,  ;,„.!  v,.n  llc'ss-  to  -nak.  ..  ^aiv- 
,•,.1  stu.lv  in  rats  ..C  ll,.'  vas.-ular  n.tl..s.s  tl.at  .m.uI.I  I..'  .U-it.-.l  fn.n. 
^a,i..Ms  i.si....s  in  tl,.'  s,.i,u,l  .-..nl.  Tw..  i<in.ls  ..C  vas.-ular  irll..x,.s  xv.'.r 
siu.li.'.l,  l.n'ss..r  Mn.l  .l.,.r.ss..r.  tl,..  l-.-rn...-  l.-in-  .•li..it...l  l.y  str„n.„' 
,,„|  ,|„.  1,,,,,,,  |,v  v.rv  tV.-M..  stinu.lati..M  ..f  tl,.'  .■.■..! -al  .^n.!  ..t  II..- 
s,.i.,ti..  ai.l  l.ia.-l.ial  u.tv.s,  TliVv  (-....n.!  tlial  llu'  patl.wavs  l..r  pivs- 
SUV  nn.l  .l..|.ivsM,r  mIT.  iTi.t  inii'uls.'s  ^^vn'  .|nit.-  .litT..f.n1.  Thus,  ait.T 
1,„,,1  l,..„,iMM.ti,.n  ,.|-  tl...  ...nl.  111.'  .l.'lM"ss....  r..|l..x  ..l.taincl  l.y  M.-aU 

stin>ul,>ti..n   ..r  tl:,.  s.-iati tin-  s,-,n>..  si.l..  as  tl>..   l.si.,n    was  n...ma1. 

v^lM.tvas  it   xvns  ..T-.atlv   r..! ■.!   uL.'n   tin-  s.-iati.-  n.-i'v  ....   tl..-  ..pp.'s.t.- 

si.l..  r.-.,...  tl,,.  l..si„n  XV  as  stin,..la1,.,l.  <•„  tl..-  -tl..-,.  l.an.l.  tl..-  pr-ssoi- 
,„„Hi,.„s  tl.,-.t  w.-f,.  .„.,st  ..,a,k..,lly  .r,.„^.,isl„,l  NV.-f.-  tln.s..  f.-,M..  11..- 
s,.i.ti,.  ,m  tl,..  sMM,.-  si,l..  as  tl„-  l.-si..n.  Tl,.-  ,l.-l..-,-ss..r  lil..-.-s  .-v.-l.-ntly 
,.,„ss  in  tl„.  ,-,.,•,!.  >vl....'.as  tl,.-  i.n.ss...-  ,1,.  s„  ,.nly  t,.  a  l,n..t.',l  dv^u''^. 
K,„.|l„.,.  it  WMs  f.,„.„l.  alt,-,-  ,-.,11i.,'-'  a.-n.ss  tl,.-  p.-st,.,'!,-!-  pa.'t  ,.f  1  ..' 
,.„,,,.  „„„  ,1,,.  p,...ss,...  .•.•II,-x..s  AV.-r.-  i..t,-.-IV...-,l  xvitl.  1...1  ....t  11,.>  .  .-- 
,„-..ss..,-  thus  i„,li,-nti„..'  Il.at  tl„-  \unuvv  a,-.-  t .■a..s,nit|...l  ...tl,.-.-  I'y  11,.- 
,,„st.-,i.„.  .■..l.....n^  ..I-  "l.il.-  n.alt..,-  ..-■  l.y  tl..-  --May  ...att.-,-  -r  tl,.-  p-st,-,-,,..- 
l,„,i,s  T,.  .l.t..|..„i>i.-  xvl,i.-l.,  .■xp..,-:.....nts  ^u'Vi'  als.,  p..||-..i',i,.-.l  in  ul.ol. 
,1,..  ,„,.t...i,.i-  .•,.l...„.is  w..,-  al..n.-  ,1.-st i-oy.-,!  a„.l  tin-  f.-snlts  .■.nn|.a..-,l 
xvitl,  ..tl,<-,s  in  «l,i.-l.  11..-  tip  ..!■  tl,.-  ,...st,-,i...-  l,..i-..  was  .n,-ln,l.-,l.  S.-.'-." 
it,  wMs  „„lv  in  tl„-  latt.-i  ..Np,...i.n.-..t  tl,a1  any  int,.rf,-.-,-...-,-  will,  p.vss,,.-  .•.- 

,|.,s,.s  was  f.„u„l  t •...•.  it  was  ,.,.„,-ln,l..,l  that  tin-  i.,.s1.-.-i,.f  h,..-n  al,.n.- 

is  .•,.n,-ii,t.,l  ii,  th,.  1ia,,siiiissi..i.  ..!'  pifssi.r  iiiipuls.-s. 

He.rnnlin .n,h.<-ti,...  .'f  th.-  alT.-f.-nt  i...p,.ls,.s  whi,-h  in  .-.ms.-i.msnrss 

p,.,.,l,..-,-  pain  a.„l  ,.!'  th.-s.-  .-.....-.-.•...-.l  i.'  tl."'  '•.■"'•>^  ''1'""*-"'^  '"'  '••■>^l""<""""- 

it  was  f.......l  that  tl,.-  post.-.i..,.  h..r..  .-i"  !-'>ay  ...atfr  is  i...t  .Mmc-.-ncl. 

fn„n  wl.i.-h  it  is  inf,....T,l  that  s..,-!,  i...p..ls,s  a.'.-  <-,...,l..<-t.-.l  l-y  tl.o  sam^- 

i,lT.-i-..iit  l.ath  that  is  inv..lv.-.l  in  th.>  d.-pf.-ss...-  r.-ll.-x;  that  is  t..  say,  as 
u,-  havo  in,li,-a1,.,l  al.„v,-,  1h.-  impulses  .•r,.ss  in  Il„-  vuvd  t,.  Ih.-  ,.pp,.s>t,. 
si,U-  an,l  as,.,.n,l  in  tin-  lali-ral  iuni,...lu..     The  pathway  ,.f  1h.-  .-p.cr.li.- 
an,l  l...,-ss..,-  s,.nsMti,...s  i..   .1..-  ••,..■.1  is  .."t   w.-ll  know.,.     It   .s  l...lu-v.-,l. 
h,.w.-v..r    that   in,pnls..s  „f  t,...,-h  pass  up  the  iv.ste..,..-  .-..hnnn  on  11..- 
sai.ie  si.le  of  ll.c  .-"i-.l  f".'  f.....'  <.>■  tiv-  s(.-.'n,.-..ts,  aii.l  th.-ii  irfa.l.u.lly  pass 

to  the  anl.-ii.n-  .'..lim.n  .'I'  the  opi).)site    i,l.-. 

But  f..r  ol.vious  r(-as,.i,s  it   is  mainly   f,-om  ,-li„i,-al   ,>l.s,-..vatlons  an-l 

a,.cu.-a1e  p,.st.u,...t..m   l,..-at,on  of  ih,'  sp I   .la...a..,.   that   th.^   pn.hU-... 

must  finallv  h.-  solv...l.     I'-y   Ih.-s.-  i...-11.".ls   it   has  l.c-u  shown   that  sni- 
s<it!o„s  of 'pain   a>i<l   l<n,i„r„t>m    pass   throuuh    'In-   opposite   latrnil   col- 


II  \iMi\    .11  liK>  :  M 


VMilO     I'MIIW  \V>    IN    >n\  M.    I  null 


:i:l 


„ ,v.    ..,/.     >■-»..     Ihnnnih    II.,     I -,/../'-•.<'    •/"'■-'/    <■"'"'"".    "•'"''     '"''''• 

,,„„,/;„,->•    /.">x    /"""•/    '"/    ""     "",■-•.,«,,/   /-/.,.■>■    „/■    //..     ./,„•>.-/    r„\,n,>o    .,/../ 

,„„-ll„    l,„    tin    .,,., '.     IxI'r.u    ruh.nn...       It    |.    iiit.n.Min..'    llud    of    tlu-,' 

two   |.;lllis    t,.r   tlirlilr    1 Ills.-.   111.'    .-n-s.Ml    nIM.    is    m1 Insrlv    ilss.  "MMf.  1 

U  Itll    ill.'    Iiai-I     lluil    1,11  I  irs    pillll    I  l|ollilr>.. 

1|,.,„|  ;,|„1  Thonii-M.,  '.,iv..  ills,,  r,,uii.l  tli;it  til.'  s,.|,s„ti.,iis  an-  ..m,,.i|.  .1 
,„  ,1„.  ..M,.n1  lliat  thos,.  „f  „!,.■  kni.l  liaxH  t..-.'tl,.T,  ulirtli.T  iIp'v  ai- 
,•,,„„   ,1,,,.,,  ,„    Mi|HTli,-ial,   I'lnin   |-r,,t,,|,,-itlii.-  ni-  rpirnti.-  IT,-..).! .il's.      W  li-ii 

,1„.   ,,,,,|.,.,.,ali r  .-Mtaii, s    |,aln    is   I„s1,   s„   mK,.   is   tliat    l.l-.„lu I    I'V 

.l....|,    i.n.ssiiiv;    liulit    lull,.! i    ioaNV    l,.u,.li    arc    als,.    l..st    siiiiiiil ■ 

,,ilsl\.      Til,'  a|,|,iv,M,iti,,ii   ,.f  all   ,l,';iiv,.s  ,,l'  |,.iMiHTii1lir,'  is  aliolisli,-,!  at  lln' 

..,„„■,  ,i Til,    ai.ililv   t,,  ilis,  riiuiiiat,-  li-'tuiM'ti   In,,   |„.ilits.   tli.-  alM-l la- 

,;„„  „!■  u,.|..l,t.  tlir  ,,',-„-iiiti..ii  „r  111.'  vil.ratioiis  ,.!•  a  li-'avv  tiiiiin-  (ovk 
.,|,|,li,.,i  I,,  til,.  sUill  all  '\r\nU.\  ,„l  il,l|,uls,s  ,.„i„luH,',l  tlii-oiiuli  III- 
Il,.lll,,l;i!i'ial    ,|o|siil    ,',,lllllllis. 

l',.,.Mi|s..  tlir  ,M-,,ssiii,j  ill  111.'  .'..r.l  .ifs.'lis.,iy  lili.'l's  ,'aii>iii.^  .•.•■tain  s.ii- 

sali..iis  ...'.-Ills  iiM.i'.'  i.r |,ll.v  than  tliat  ..f  iIm-,'  .-airvin..'  ..lli.Ts,  an,l   l',.!' 

„,1„.,'|,...  ,.l..M,'lv  mi.l.'l'sl I  l'.':i  o'l^    ll Iini.-al  r,ii,lii„.'s  m1'.'  ..ft.'M  .lilli.-illt 

„,•  i„!.',-,,i'..tali..n.  ,'sp,..-,all>  u  L.'ii  tli.'  I.'sl.ais  an  ..ul.v  nai'lial,  Tl." 
s,.,is,.s  .,r  i.aiii  an,l  t.'iii|M'ralm'.'  ar,'  iiii.l..uM.'.ll.v  List  iini.-li  in.,.'..  I'.'a.lilv 
tliiin  lli.is.-  .,)•  .•iilaii.-.uis  s.,isil,ilit.v.  lliou.^'li  soin.'liiii.'s  tli,-  r-'v.Ts,.  ,-,.n- 
.litio.is  an<  found.  If  a  partial  losiou  of  .Mie-half  of  tl:.>  .oi-.l  o.vi.rs 
nimnt  the  level  of  the  twelftli  dorsal  sesinient,  n  very  eommoii  sym,.t.mi 
is  loss  of  powiT  niul  temperature  on  the  opposite  side,  l-ut  not  ..f  lou<-h 
oven  ulien  stn.iiir  stimuli  are  applied.     This  .•i-osse.l  lelation  -Iocs  not. 

),„w,.v.'i-. ur  "li.^ii  the  l.^sioii  is  i,..|.iu   lli.-  Iw.'llUi  .l..rs:il. 

I,', '-.'ir.liii'-'  111.  Hunilx-r  of  sc-^iii.iib.  ii.  .■.•ssary  for  tli.'  .l.'.-nssali,.ii  .'t 
.'ii.-h  kin.l  ..f  s.'Ms..  til,.':-.  oli-fr\  aTh.-is  on  .-asi's  in  ulii.'li  tli-'r.'  is  iiiiilat- 
,,,,,1  ;„i,M'v  of  III.,  .'..r.l  aiv  l.-'ini.'  .-oll.vt.'.l.  -.  tlal  tli.'  i.pp.T  lii.iil  ..f  Hi" 
niM'sthi'li.'  ar.'a  ina>  I..'  .'..nipar.'.l  with  th.'  si'LHio'iilal  l.'V.'l  ..f  'h,'  injurv. 
It  a-.,„,i,'s  thai  IKiin  an.l  tla-niial  lni|.il!s,'s  ,'r,'ss  ,|irH'U!.\  ^  i.  .'..  uithili  a 
s.Mnm'ii'  or  t«o  in  iIh'  i,rhl.l..rsal  r.-ion.  luit  that  llms,.  of  I..U"h  .•!•., ,s 
soiiH'uhat  Mior.'  .^ra.lnallv  In  tli.'  upp.'r  sc-.^miiits  the  ol.li.p.ily  of 
,.,„ssiin.   of   iHiih    kiii.ls  ..f  tilu'i-s   is   ur.'at.'r.   aii,l    in    tlu'   .-.'rvi.'al    r..-ion 

i,    ,„;,v   ,' ir..   ti\..  or  six   s.'iiiii.'iils   for  the  .•r..ssiii..    of   pain    inipuls.'s. 

With  this  in.'i-.'asin.^  ohli.iuily,  a  .lisliii.-tion  a|.p.'ars  ,  lli..  .•rossinM;  l.-v.'ls 
„,'  ,,.,in  ;,„.!  1,.|,i|M'ralnr.'.  for  ih,'  lalfr  ,'ro>s  a  littl.'  mor.'  .piiekly. 
This  .■oiif..nns  uilh  Ih.'  .-Imiral  ol.s.rvali.^n  that  llnriiial  appuM-inlion 
,„av  he  .listurli.'.l  wilh..nt  lii,-it  of  l-ain.  Kv.ii  th.'  lln'mial  iiupiilsrs  .).. 
„.,t'  all  de.M.ssate  af  th."  sani.'  h'v.''.  for  an.'sth.'sia  I.,  h.vit  may  n'a.'h 
hi..difr  up  .111  th.'  skin  area   than   lli.'il   to  ■•ol.l. 

■\Vh.'n  r.'.-.iv.'ry urs.  th.'  s,.iis,',tions  -^ra.lnally  ivappear  .•au.lalwar.ls. 


f^l 


J 


I 


ill 
1      : 

ii. 


jy 


H 


"^iii 


I'll!     (  I  N  lltM      M  U\iil  >    ^V^^I.M 


Solll.lilll.H    III     lli'-'ll     lrsiMl|>    nl-    till r.l     lIl.'IV    i>    Jllicstll.si.l     Mt     I  lie    cnr- 

IvslmlHlill-    l.V.l,    hut    III.'  lU.M    Mippli.'.l    I..V    llir    I..U.T   >|.inill    1 IS   vs\„: 

rinlly  111.'  ^Kin   in   Hi.'   I'-uioii   i.f  lli.'   mmu>.   is  ><'u>\U\>'  \><  vu •  "tli.T 

UilHl     of     MilMlll.-ltiun  III      IT.-nV.TV.     tu,.,     tll.TC     hlllV      \<r     iHI      .Jlllv     IViip- 

,„,;,n,iii'..  ol'  vniMili-.n,   in   is,,lnl.-.l  .•ini.liil  iin'.-is.     Tin'  .■sphiiiMl  t..ii   -iv.M 
r,,,-  lli.s,.  iv>iilts  is  lliat    III.'  lilx  r-  r.inyiii-  .lilV-'iciit    kiii.U  '<<  s,.|is;i1  i.'ii 

ii;iv.'  a  hiiiirlliu'  iin'aiiL'.'iii.'nt   ii,  tli .1.  lln'  Imi-.'st   IHmts  Im^Iim,'  ..ii  IIh' 

oiitsi.l,.    .MM.  |M-.'  M:!'.     NVh.ii  .'1    iMitial    i"si"ii   .'ilV ■<  til.'  iii.'sial   lili.'i's 

„„,iv  lliJin  till'  laliial.  tli-ii'  "ill  nr.-nnlin.J.v    I-  i-.tux.iv    m'  l|i lUilal 

-kill   areas  licri.rc  tliosc   liiuluT  np 


i    ! 

j 

, 

■ 

1'' 

J, 

\^^ 

:  « 

i  r 


^H| 

M 

H 

fi: 

■^  i 

I' 

ill  IT: 


CIIAI'TKIf  X'lV 

KiiKtis  oi' i;\i'i:i;iMi:\T.\i,  i.i:sio\s  ui'  \.\i{i«>is  i-auts 
(ir  Tin;  nkuvois  svstkm 

ll.-i\iiii:  Iciinid  till'  main  .'li:!!  iic-l(iisllr>  „\   till<  \  .-H-liMii,  "  c  sliall  H"" 

|l| ,v.|     til    sIllil.V      llll'     prrlllKir     riHM-tinll     .  i  T    I'llch     jMl't     of    til.'    ,•,■  Ivl  1 1'l  in|  .1 1 1  ill 

s\-.|rlll     liy     lli.tilli;     til.'    I'lV.ct^     «hirll      lujlnu     .l.^^tlllcti I'    -t  i  lllllla  I  i.HI     "!' 

'its  ijirrri'i'iii  |iaiis. 


THE  ANTERIOR  ROOT 

Scclinii  (,r  all  aiili'iiHi'  iii.it  |ir.M|iH'cs  a  liiiiiti'.l  .ic.j-ii'i'  (if  paialvsis  !)!■ 
Vdhiti!,'  SIMM-ill  iiiils.-li>  liji\iiiK  HI.  Iiiiii'tiiiiial  iclatiuiisiiiiis  111  mil'  an 
otlicr.      If   si'vcral   aiilirini-    imits   air   i-iil.   tlif    paralvsis    Ih'I'iiiih's    imii-li 

limn'  cNti'inli'il.  iiml   is  t'nlluw  nl  vii*    s i  hy  iiii  i'\  iilfiit   iitinpliy  nf  tlif 

niiisi'iis  iMiiii'iMiiiil.  Iiillis  iiclii.iis  fnnii  tlicsi'  iiiiisi-jfs  iiic  "(  i-ihiim'  nn- 
iiiissililc,  Stiiiiiiliiliiiii  111'  the  |i.'iipliiral  fiiil  nl'  a  ciil  iiiiitnf  rmil  ciiiisis 
]iaitial  riiiiliiii-tiiiii  of  si\cial  iiiiisi-lfs.  im  liclinilc  jniiit    iiiiiM'iiii'iit.  Imw- 

I'vi'i-.  liriiiL'  till'  ivsiilt.  I HUM'  till'  alVcrl.'.l  nnis.-li-s  iiir  nut   li timiiilly  iv- 

l.'iti'il  iiii'l  tliiiT  is  nil  ri'i-iinural  inliiliil  iun,  I'Ii'Mh-  ami  i  stiiisor,  inliliii'lor 
ami  iilMlm-tur  may  iMnitriu-t  at  tlir  saiiir  tiim',  thus  caiisin^'  tin'  juiiit  mi  wliii'li 

l|i,.y    ,Mct    til   111 nil'    iiMisi-li'liiiiinil.      It    is    ill   till-    iili'xus    that    thi'    iiitvc 

lihiTs  lit'  till'  iiiiits  iM'i-iiiiir  Miiliil  uiit.  iiri-iiiiiiiiu  tu  I'liin-tiuii.  intn  iimtiir 
iiiiil  scnsiiry   ihtvi'  trunks.     The  ilistriliiitimi   ut'  tli.'  antiTiur   runt    lilifi-s 

iiciMirilini.'  til  sriiiiii'iits  in  iniii   fur  tli i\ii';il  iiinl  liniilii.siiriiil   i-i'>?ioiis 

is  iis   fiilliiws: 

(•-,        |l,l|Mi.|,   Lin  |iS,    lilil.  Ili;ili>,   Ml|iiliil  ■!-.    ill. 'lilt... i.N.      I)..;i-i \\\ 

lii.liiil  .■\l.ii-..r-J.     l.'iiii-ty  |.r.in;il..r  ni.lii  lirc^. 
Ci!        I'i..ii.-i|..r-',   '         .1  lAl.'ii'.irs,  pcctnriilis  iii:ij.>r    (.'liiv  i.-'il;ir   lili.Ms~i, 

s.'ri'iitii-  ;iiil  i.'its. 
('7       •'l'ii.-.'|i-,   .'\t.'iis..i    ."ii|i    iilii;iii~.   .Alciis.i's   ..r    liii;;.is.    |..-.'l.'i:ilis 

iii.-iji.r. 
Cs        ri.  \..is  .if  »i  i>l  1.I..1  lii,:;.'is. 
'I't         Iiiti  iii'i.'  niii'-.'l.'s  rif  |i;iii.| 

s:;.  1    I.inul.'i    ;i;.i,  >|.|.iii.  I.'i    iii.i.    |i'iiiiii.l    luus.'li?. 
S-J        (il.il.'i,  l.ir.'i.s,  v.init.'ii.lin.isiis  iiii.l  .-. '1111111. ■iiilir;iii..siis. 
yl         !.:t.;!i-;.-  f"i-!.-  ■■!■  f.-.t.  !;hi:.!U  ju^-H.-is    ;.i..!   !^r:<Ar<   ■":  ■■■.:\>:. 
I.:,        Miis.-lcs  of  viiilniliilCKil  1.;;   (.s.-.'l't  lil'iiilU  iiiili.-ii-.  i . 
I.l        i:xtiiis(iis  of  li'j;  an. I  til.iiili;  nnti.-iis. 

s:;.-. 


i-  ii 


•»:!(; 


nil    1  I  s  111  M    M  mill  ^  ■^v^ll^l 


t? 


ir: 


m" 


1  •    !. 
i 


A 


■;i| 

'  •  it 


AV   if 


i      i 


I 


d'   IS 


Tlir    klH.ul.-.lu.'    "I    111"    •..•um.lili.l    IMlli-.tl    •'(•   llir    Imili    liill>.-l.'s.    i,. 

nni,i,lir.l    ill    tlir   .iIm.v.-   Inl.lr.   is   ol    Mihir    in    tl.-   I.MMJi/iil  nm    uf   si-iimI 
l,.Moi,s       l\M.il>Ms  nl    III.'  .M.n^i 11. .Mill,  hi,  ..!■  III.'   uiisl    ;u,.l   lillK.TV 

;,|,„,H      Ullll     III.-     Ill.'.|.v      r..t      .AilMl|.l.'.      IIMIilllN       in.ll.Ml.^     M      I'-' t'     •'"■ 

..^.,„lli    ,-.iM.al        It     i>    V    |...lll.-iihiil>     ill    III.     iMlliU,    l,..u.\.T.    iIkiI 

t|„.   s,..iii.Mlal    iiHi.Mv.ilMUi    ..r   til.'    iiiiis.-i.-    i>   .'vi.l.'iil.      Til.-    iiili.i-viili..n 

„(     ,|„.      i,,t.T,-nMill      lim>.-l.->     ll.-ll'U      Hlls.-L'.ll.-lllJll.     ..II.'      linl>       .ll.l.^lM.v.-      IIh- 

|,.\,.l  .,1  ;i  l.'>|..ii  ..!'  111.'  iil'l"'!'  Ili"lit'-"-  l''i;i""  "•'  III''  '■'"■'I  ''>   "•'-'■'■^iiil-'  ''"'"■ 

l„.i,,,vi..i-  .liniiiu   .i.-.-|>   ins'ii'i>i"ii       ll    ''"•   '■'"'-"■'■"   •"■•■    !''■"'"'    '"    " 

,.,, Mill    >|.i..'.->.    til.-    |.;ii,ilv/.-.l    iiiii-l.-    uill    t.-l    li-ip   ■•iii'l    •'"•    l'""-"'^ 

-ink    int..  til.'   >|>a.'.-  .hii  iH'-'   tli''  .I'-l 

|.,„-;,li,,,li..ti     iii.i.v     .ll-.     I'.'    «li""ii     l'>     -'ii'lvii'!     '!"•     I'ai.|l>v-   .'I-     III.- 

,,|„l,,i,,ii,;il  inii--l.'-  "Ih'Ii  til.-  I.-M..II   ii.'.i.iv "\   111''  I""'-'-  •'i''  tl""'i|'''"' 

s, -1,1.  Ill-      Wli.ii  111.-  I.ati.-iil   uitli  .-I  l.-M I   111.'  .■I.'\.tilll  Ih.-nt.'i.'  I'Mi  — 

In"   I, .-M.I    II. .ill    111.-   I..''l    ..1-   i:li-.    Ill''    '■•''•'"■^   .•..iiliii.-lv    I. ill    111.'    ilNH'    ''' 

.jh,n-  l,i.l-.-..uili'„'l..  |.:iiiil>>i>..rili.-  I..«.'i'  |..'rti..iiM.l'  III-'  .■l.li.|i"'--     I   "'I''' 

,|„.  sai |ili,.|iv  uli.'M  III.-  niiitli  s.-^iii.  lit   I'  \nU'Ur,\  111.'  I'.-.'lil-  .■..iili'a.'t- 

,,,  ,„  ;,|„,|,.  ,,|,.'  i.i.-li  ;il'..\.'  III.'  innl.iri.'iiv  wli.'r.'ii>  l..'l..«  ll'l-  I' ^''l  ''  >'''lii''ll"- 
iliii-..|ill';i.-l.-.l.  -n  'lial   ill.'  mill. ill. •ii>  i->  |-illl.-.l   II!'. 

|'.,.,i.l.-.  iiiii>.'iilai-  iii..s.'ni.-nl.  >rniiiihit  l.ii  ..f  iIh'  anl.'ii"!'  i""ts  in  li-litly 

,in..stli,-ti/.'.l    iii.iiiuls    M.in.'ti.M.-s    .',       .'^    .vi'l'' '    "!■    •-■'■ii''t''i'    '■'■"'"^    '•'• 

s|,„M^.'  :iii.l  ..f  l-.-iiii      Tl M.liiiKiti..ii  i>  tliMl  tli.'i'.'  .'II.'  I'l' lit  in  111''  "I' 

,,,,„,  ,  „„,  ,.,,,,.,i„  M'lisni'v  tili.'i'^  u  lii.'li  Ml-.'  .l.-iiM-.l  11  ..Ml  111.'  |...-<l.-il..r  r....l 
|,„t  I'.'.'iii-  in  ll  ■  iiiit.ii..i-.  -  11^  t..  |-.;,.'li  111.'  ni.-n,l.nin.'>  ..f  lli.'  S;in'il 
,.,n',|   vvli.'i-.-  tll"■^    l.-l-liiilnit.'.     Til.'  slinilll.Hi.'ll  "I'  111''  I"  "i'''    ■     -li'l'-l'"' 

„,„,,,,  ,',,., 1   I,, .1-1    |.1'...1m.'.'.  tli.'1'.'r..r.'.  111.'  -a r.'ll-'N   t.'M -'-  ■'-  ^'""" 

i,,ti.,ii  ..r  111.'  .'.'iitnil  .'11.1  "I'  111.'  M-1I-..1V  r..nt.  Stiniiihili..n  ..f  Hi.'  '"'H 
,,..,]  ,.|,,|  ,,r  :i   in. .!..!'  i-....t   li;!-^  '"■  .'"III--.'  11..  .-IT.-.'I. 

THE  POSTERIOR  ROOT 

TIh-  ,,„v^  ,-;..,'  ro„l  i,  1 1..-  |.ath»;iv  \n  ulii.'li  i.iil.uls.-s  .-I'  tl..'  Vi.l'i..ils 
,,,,..,„.„'.  .'iit.-i'  III.-  -filial  .'..nl.  S,-.'ti..,i  ..f  anv  .'..nsi.l.'ial.h-  numlM-r  ..i 
,,„st.'i-i..i-   i-.M.N   .-an-.'-   tli.'n'lni-,-.   ai,.'slli.->ia    ..1    Hi.'   .'..rn'Mi..n'lHi--'   ^'  i" 

,„„1   niiis.'l.'  a.'.'as.  In.l   Mi.-l,  a   n-ult   .|.--  iml   1' ' vi.l.'i.t    u  li.'.i  -.i-' 

,„„,  al..n.'  i-  .'111.  iM'.'ails.'  111.'  •^.'n-.-i>  n'''-'  >H|.|.li'''l  I'.v  -'a.-h  l'....t  nv.'i'- 
).„,-  ;,t  l.'asi  lialf  ..r  that  suiipli-'-l  I'.v  Hi-'  ti.'iL'lil...i  lui:'  i'-"ts.  Altli..i.'Ji 
•„  is  oi'iin  .lilTi.'iilt  t..  -li-tii-A-ii-l'  ll"'  -<"^'ii"i"'-'l  -li^tiil'iiti""  I"  ""■ 
,.,,nii;.atl..n  .if  tin'  IH-.-'h  "f  Hi.'  nn.t.u'  mots  I.v  Hiulin-  uhat  .ni.s.-l.'- 
tli.'V  intln.ii.-.',  tlii-  is  ..MM'.-  .'vi.l.  nt  in  tl...  .-as,,  of  tin-  M'nsory  m-ts.  On 
,1,..'  trunk    it-.'lf  tills  s.'uiii.'iital    an'an-.'.'nn'nt    is    vry    plain,   l.nt    in    Hi.' 


i'i   ll      ! 

m 


I  KH  I  r-  "K  I  \i'i  unii  M  \i.  i.i»iii\!- 


KIT 


.•MnMnli.>  II    l>  IM.I   at   li.-l   >iulil   su  ••l."'.  mIi1m.i|..'Ii   H   -mh  It  i iiral-lv 

WnlUr.l    ..l,t.   Il>    is    ll.dl.Mlr.l    til    III.'    II I|IIIII.V'I1..'   .lui'.-nilll    <V\'^      ■1\^ 

III  i.ii.'iii|.iiiii.'  t"  .i.'i.'iiMiiH'  III"  i"^''i  -r  II  i''-i'"i  I''""'  ''"•  --ii-"''.^  i"""'> 

si,,    sn. ,1 -I'll    -III-' ""    "'■    '• '''"I" ■-"    "''    ""■ 

,|.v,...i.ti !■  III.'  -ii-.i-v    til"'-  III  IIm.  .|Mn,,l  -•..i-.l.  lilMr>  !,.,■  tl  ■•  .lilV.ni.l 

s..Ms,,li..ii>  noi  .■.■n>.in..'  Ill   III-  Slim.'  I.-..'I>      l'"i'  .Aiiiiirl.'.  Hi-  ii|'|" "'• 

„,■  „„„|„,..„,.  iniip.  iiiliii.'  is  "It'll  l"-l   sli'^liil>    liii^t"'-  lli'iii  '"""   "'■  I"'""' 
.,.,„.   „,,,„,...,MliuM    Ml    ilir    mKiiiIimi,.   .ails.. I    h    •li.nMiiU   Hi-'    I'n--'   "''   ^i 


\.ti    -i-     1' 


1,,.,,,      \-: 


lu.avA  tuiiiht:  U'vU  ..v.r  llir  skin  is  oit.-,i  vny  iisHuI  in  l.-rMtini;  tlio 
h.sinn.  iiaWi.-nhirlv  in  Hi-  al»ln„un.  Wli-n  lliis  iii.-,1,.m|  is  usr.t  on  ill- 
tln.niN,  h..urv-r.  ill-  skin  s|„„iM  l.r  pull-.l  .i|,  in  r..l.ls  l-Hnr,.  ill-  fnrk 
is  appli-il.  siiHM.  ,.lli.rN,is,.  111.-  lli.Hax  uill  a.-t  as  a  i-s..nat..r  ami  s|..va.l 
tl„.  srnsati..n.  S,..-li..n  ..f  tw.i  or  iii.'i'.'  s.-ns..r.v  i-.m.Is  i-r-Mln-.'s  a  very 
,,,.|i„i,,.  ,,,,,,  .,,•  anrslli.sia.  inv..lvin..'  all  lli-  skin  s.-nsa1i..iis  as  «-ll  as 
several  of  llmse  of  .l-.'p  scnsali.in. 

llier-  is  not  only  an  .ntirr  al.s.'.i.-  ..f  sensati..n.  l.ut  a  niarke.l  intertVlTl- 


1 


}.,rM 


Mi:;l 


I 
i  ' 


■ti.f 


j;..jj  Till      rlMi;\l.    MUM'I-    SV^TIM 

„i,l,  ,1„.  nv.ruln.-  or  tl,r  I.Mil..  Ilr  roiMl.linn  Wuvj  .Mll.'.l  ^r/„s//M.,„. 
Tl„.  ..xa.^l  ivMiU^  .l.lwiHl  MMiH^x^ii.M  on  r„r  lv|„.  ..f  nnnual.  Il  all  11m' 
,,„sl,.|ior    ,o„N    ul     llir    :,nto,i„r    rMrnnily    aiv    .-111     ni    .-i     „o-»/."/.    tin 

.,,.., lin-  liMil.  ^^ill   nm    1-   nsc.l   in   .-InMlnn^  or   l..r  ..Uirr  ,.„r|."s- 

I,     „i||    a,,,..,-.r    In    I iil-lrlrls     paralv /.•.I,    unl.--    ^^hrn    tl |.|.os,l,. 

,„„„„,      |i,„l,     i.     ni      Ni.n.nus     a.-rui,>.     ull-M,     Hi.'     Mposllutlr     .-MM,     UniV     l„ 

liiuM.I     111    ;ivMiri;ition. 

Or.   r,-i.vhil   rxniniiutinn.   Imu  .mt.   il    «ill    1"'    inuihi   Hint    iinirk.'.l   -lil 
,■,,..„.■...   oNisI    in    l!,r    IVI"-   of    l.iialvsi.    |,ro,|u,.r,|    Lv    ll,.'    srHiol,    ol    11.' 
,,„,,,•„,,  „,l  tl,..   post.n.n-  rout-,.     Winn  a   uu.un-  mo,    i.  ,-iit    no  nllrxr. 
.,,-,.   pos.iKlr   ritiior   tVoiii   tlir   skin   or   (ynm    tlir   ,Tirl„al   .•oitrx,   ana   tiir 
,unM-l..s  iiimIci-o  ali-opliv.      Alt.-r  M-rtio,,   of  tl,,.    |.os,,.i-io,-   ,•,:„:,  o,,   tl,.. 

nth,. I-    lian.l,   allln.uuii    iHlrx...    Ii- !!.■■    ski,,    ann    ■A((rr\.A    arc    impos. 

mM...    v.!    inovrlii.-nts    may    !.,•    .■li,Mtr,l    Ky    aitilirial    sliMiillat  loll    ,'1     tin' 

,.,.,-rl',ral  coi-t.-x,  aii.l  tl,c  iiiiis,-!. -  ,1 t  ali-..pliy  to  llir  s,-,nir  .•xtont. 

II  ,,nlv  on,.  >,  nsnrv  n„,l  of  a,,  .■xtrmiity  is  I, -ft  in, cut  for  cxaiiiplr. 
,1„,  |,,.|',..Tvical  s,,  tliat  tl,r  skin  of  ti,r  !,ali,l  is  slill  snpplinl  VMtll 
srnsatioi,   knt   all   tlio  ,k-P   i—rpto,.  aiv    -v.'iv.l.   lli.i,   li,,'   liiiil>   may   1- 

nM.I   to  a   I lilir.i  .1,  uiv,..      It    iiia.v    Ko  iis.mI   l.y   l!io  iiionkoy   to  j.n-k   up 

,.„|s  Init  tl„.  iiioNriiin.t  uiil  1...  liistiniMly  chinisy  an.l  ataxi.-  in  natiiiv. 
Instrail  of  nratly  pi,-kiinj  ii|.  tli.^  nnts.  !„•  xvill  mak,'  uiM  laov.iinnls 
illiil   oftc,   iniss   tiiclll. 

T!„.  apcslli.'sia  is  n,,t  s,,  ,Mofo,in,!  i,,  lo«,r  animals.  Aftrr  sr.'tioii  ol 
all  tl,r  s,.|,s,,.v  v,,o1s  to  l.olli  l,i,,.|  liiiilis  in  thr  ./■■■/.  tlH'iv  'iia\  1...  a 
i,,,,,i„  attnai'.t  at  valkiir^  on  tl,r  part  of  tin-  alT.vtr,!  iiialo.  that  is  m 
sav.  ^^ll.■^  ll„.  animal  tri.s  to  pro.jr.ss.  ,1,,.  kiml  liml.s.  allli.m'J,  at  lirsl 
,„,.,-..|\  lira;;-.-.!  akm-  lli,'  -io,ni.l.  a  ft.'iu  anls  Im'-Iii  t,.  cxim-iiIo  walking 
inuvcimiits,  ul,i,.|i  hn^^,■^,■r  aiv  v.n  jrrky  or  ataxi.-  in  naliliv  an.l  cMi- 
ii.jl„„,.  linl..  t..  tl,.'  |-..ruar.l  |n-,..j|-..ssi„n  of  tlic  animal.  altl,o,ioli  1,. 
inav  sii.-.-.-.l  t..  a  .■.■rtaiii  .■xl.'iil   in  siipp.nl.n-  tli.'  Im.ly  l.y  ik"  kin.l  limk- 

Tl,..    iM,p..rtai,. f  tl,.'  s,.ns.,ry    n.ot    in   .-out  ,.,l!i,i-   tl,,'   .■..lit  ra.'t  ion    oi 

II,..  IIIUS.-1..S  is  fnrlli.r  illiis|,at,..l  ky  .•..nipariio.;  tl,..  .-.mt  ra.'ti urv."  ol 

a  niiis.'l.-  i.r.,.lm-.-.l  kv  stiiiinlat  iii'^  il-  n,,..ut  i,i..t.'n.  n..rv..  with  tkat  i„'.i- 
,l,i.-...l  kv  sti,„iilatin..i  tl„.  p..|ipli.ial  ..ml  of  tl,..  .Mit  n,.|A...  In  tlic  form.., 
,,,.,..  ,1,..  .-nrv..  is  „,o,...  pr.,l.,n.j...l  an.l  sl,„us  a  -la.liial  n-laxat,on 
wk,.r.Ms  uli..n  tl,.'  p..rii.l„.ral  on.l  ..f  tl,..  .-ill  iltw  is  stiiiinlat,..!,  tl,.-  ...n, 
1,.a..ti..n    is   l.ri.f   ai,.l    tl,..    n.laxatio,,    is    f,,|k,u...l    k>    a    .iistin..t    .■.■k.nin.l 


,,r   ••in.iiia    su  ii,'j       a-    ,t    is   .-all..' 


■I'hat    tins    i|ill..|'..n.-..   .|.p.'ii.|s 


atr..r..|,t     iiMl.iiN.s    is    in.li.-al.-.l    ky    tl,..    la.-t     tkat.    aft.r    s....ri..n    ..f    tl,.' 

,„,,,, .,.i,,r  I Is,  stii„iilali..n  .,f  ik.'  iiinul    n.rvc  in  tl,..  liiiik  will  proilu.-.' 

111.,  saiii..  ..nVct    as  o.-.Mirs  »  k..n   tii.'  <-iil    n..|\..   is  si  i,„iii,it.  <'       'I'ki's.'   re 
suits  can   1...   v..ry  .•k'ail.v    ..ktain.'.i    in    tk..   .-as '   tk..    fr.        In    wki.'k 


I  III' 


ij'    1  \n.l;l\li.N  I'M.    M  ■'InN- 


839 


,,,.,,  I,  ,,111  |„,  ,,,,,,,1  ,l,;it  :,||,r  Mdi-ii  .11  tin'  postrrioi-  nmls  ,.r  on,,  si.lt-. 
,1„.  ..onvspolHlin-  lillll.  li^lML-  Inu.T  iImh  its  M-llow  lH..ausr  its  mnsrUs 
;irr    tlillllrss. 

Sliiniiliition  ..f  liir  .M-ntinl  n.l  nf  .  .-ut  .•ilTMnil  mot  i.ro.lucvs  as  l,ns 
aliT.-idv  I..VII  ih.lica!,'.!,  a  ,-.,htra-1io,i  oT  tlir  ii,ii--l,.s  ii.Toini'Min.',!  I.y  a 
iv.-ipvoral    ihliilntioii    of   th.ir   ,-,    .^    -msI^,   ^o   tliat    so.nc   .iHinitc    .n..V'- 

nirnt   of  tl,..    joint    tak.'s   |,1      -,      Tin-   •■...^ ■lit    is.  ImwrvcT,   mcirly  a 

t|,.xi„n  ..!■  rMrn>ion  or  rota  .n.  Lnl  ui'!,  n  >-yy  cvi-lcnt  oLjccI  in  view. 
In  tl.is  r.-ai<l  it  is  ,|nit..  .li  ^  t  i.  mi.  (1,.  ,.ui|.osrinl  niovcnnMit  winch 
n.snlls  IVoni  sliinnlalion  of  „  sl.i^  .■■.....  incli.-atin-.  tii-'ivron',  that  lh.' 
,,.,.,. ,,!,„■  aiMiaratns  itsrlT  iniist  .-ontrilHitr  In  th,'  ihtvc  i.nii.uls,-  sonic- 
tliiii-  whirh  raiisrs  it  t..  liiiiiL'  al.out  a  n.oir  pel  lV,-lly  intciii-at,-!  niovo- 
„„,„t  ,,!■  th,.  niuscnlatnr..  than  is  ti,c  .Msr  ^^\^<^,  the  n.Tvc  tnink  is  <ll- 
rcctU    stiiiinlatcd. 

|'.,wi,]rs  tlo.  niovriiirnts  ,,f  tlir  nnismlal  uiv  inncrvatcl  iVoiii  soirmmts 
N\hic1i  aiv  li.-idr  tiioM.  of  Ihr  stininlatr,!  alTncnt  root,  there  is  a  -.'Cin'val 
rHlcN    iTspons..  thiondi  otlirr  ,-,.nlc.rs.   lor  ,.xainpl.',   th,'   irspiralory  and 

,1,„  v,-|s tor:  an,!,  in  animals  xvlih'li  a.v  not  .l-'.^nly  ancsth-'ti/.r,),  lln'iv 

is  als,,  ,.vi,l,.n,M.  ol  |.ain.  Stinnilati,m  ,,r  ti„'  iH^rii-ln'ral  .'11,1  ,.f  tin-  s,.n- 
>„n\-  i-,iiit  has  of  ,'iiuis,.  n,)  i'ff,<-t. 


THE  SPINAL  CORD  AND  BRAIN  STEM 

Th,.  ivMill.  ,,f  lrai,^s,.,.ti,,n  ,,f  ih r.l  h■A^,•  h-.-n  ali,'a,ly  .nfli,'i.'ntiy  ,1.^ 


si'rilii',l     It   r, mains  lo  ,|iv,-iw>  t 


1 n',',t    if  total  ablation  or  removal  of 


portions  of  the  cord.  A.  «oul,l  he  .■nim.,-i..,1,  ih,.iv  i^  a  niark,.l  ,le..nv,^ 
„r  shock  for  -,i.i,.  u,',.ks  aft-'r  ahhili.m.  Diirin-  tliis  sl,o,-k  th,-  ton.' 
„f  th,'  sphin,  t.'is  an,l  vrss,ls  is  -n'atly  .l.'Pi','ss..,l.  so  that  .-ontr.^stion 
.,,,,1  ,.,h.,n,-i  ,>f  the  feet,  .li.arrlH.-i  an,l  ivt-'iitio  nrina'  aiv  n.ark.'d.  an.l 
„,,.,.,..,, i,„,  ,„■  il„.  skin  is  pra.'ti,'ally  nnav,M,l,'il>l,.,  Afl,r  a  u-x  we.'ks. 
|„,wev,'r,  r,.eov,rv  l„M.,nu.s  .-v  i.h'iit  in  s„  far  as  tin'  l,],,,,,!  x,.ss,.ls  an,l 
sphiiM't.Ts  ,an.  ,',,n,'.'rn..,l.  hut  th,.  sk,.|..t,'il  nii|s,-nlatnr,.  ,-,troi.lii..s  verv 
,.xt..nsiv,.h  an,l  <',.n.,.s  to  r,.s,.,Ml.l,.  ,-onn,.elive  tissn,..  If  th,'  spinal 
,.il,Iati,in    ii!x,,lv,s    Ih,.    th.u-a.'h-    re-ion,     f,.r    ,.xaniple.    th..    alt,.et,Ml    in- 

1,.r,-ostal    nins,-l,.s   I >nH.   stiff   an.l    |.ar,-hni,.nt-lik,. ;    the   Imnes   als,.   izet 

l,,j,,l,,.   an.l    viMl.l,.    pers,.in.ti,.n    .'an    n..1    h.'    |.r..,ln,-.  ,l.      <  Ml    the    ,ithor 

luin.k  aft,.r  s tini,.   th,'  sphin.!..|s   fun,'n..iiat..   ni,.r.'  ..r  h.ss   ii,.nMally. 

,|„.   lK,ir   is  sh,.,l   an.l   ivn,.«.'.l   in    n..rnia!    fasl,i.,n.   an.l   th.'  .applieatL-n   ,,l 
,.„|,1    ,.,    th,'    skin    .■ans,.s    tl„.    usual    sa-niar    i  ..:;..ti..n.      it    is    of    mt.'ivsl 
,,,,,,     1,^    |.,,,„.,i,.    ,,niMials    xvhos,.    Innii.ar    spinal    ,',.r,l    has    l,.vn    r,.iu..v,..l. 
ur..-,,an,.y   nia\    tak,"   i^l.-ie,'  n..;inall>,   r<.llo«.,l  hy   la,'tati,.n. 
Section  Just  Above  the  Medulla.     After  su,.ii  an  operation,  the  ani- 


I.  ■ 

I 

II 


^l^l 


i        i 

!         f 


ll 


■«i(i 


•nil      I  I  NlHAl.    M.liVors    SY>T1M 


n..M.>  ,lin„  is  ,.o»,M..  ulu.M  tlu.  s,..-tio,>  is  l.H.....n  llw  u,...lMil.     n.        h 

.,,i„.l   •.!.      Its   n.,l,.x    r..s,.nns,.s  M.-,.  MM.n.  luoa.llv    i,„...n,....l,   but     Ih' 

...„v„u.„.s  =,.-  iiuMpaM..  ..r  .■N.ruli,,^  .MovnnnHs  ,1,1,1  .,t  m1  ;„,y  val,,- 
i„  ,,,„„ns.i„„.     Mnv,.,„r„,s  lik.  ,l,<is-  c.r  ,„oon.sMu„  ,„a,v  o.MM.r,  l.ut  tin- 

„ n„.  „,HV,v:,v..,     S,„-l,  a„in,Mis  slmw  .nark,.!  s,.,...riu>M.y  o^..,•  s,v,.lly 

.,„„al   .H„.s„„  .• ,n,t    ,„■  th.  tVt    thai    i„  tl,..   .n,..l,.lia   a,v  In.-a.r.l   so 

,;,,„v  .„•  ,1,..  „n,.or,au,  .M.„„.>-s  wl,i.-l>  .-..t-ol  ..i,-.M,la,ion.  ,-..s,,„.a„o„ 
.„,.,•„„,„„,,,„,,„,„„•,„.>,„•,!„.  l.o,hMlia,  i^.  .iH'  .a-l,au.sn,s  ul„,-l, 
,.„.,.„m|.all\    tlM'   lils,    sta-.'S   i„   Hi.'  .ii-.-t,..,,   o,    Inn,!. 

Section  Just  Behind  the  Posterior  Corpora  Quadngemina     A   ^rvy 
,|is,i„.-t   i„„,.ov..,„..„,  lM.,..,n,..s  „.„i.M.al.l..  i„  111.'  ,vs,.o„sos  ol   ,1,..  aniu.al. 

Tl,,-  ,ail,o-,   has  l„r„   s,„.li,.,l  ,„os,    .•an.lull.v   i„   1i,.'  .-as tl,.'   In,., 

„,,i,.,,  ,rt,r  s,...i,  a  s,.,-,io„  ,-a„  ualk.  s,„i>,^  a„.l  su  in,  a,.,.an.n,  y  hk.  a 
„„n„al  a,m„al.  a,,,!  -i-aks  .1,,.,,  ,1,..  si,!,  of  ,!,.•  l-.ly  ,s  strok...!  In 
„,„,„a„„„alasi„„la,„HMvas,.i„,l,..-ou„,l.xit>  n,'  „,ov  ,w„l  ,s  ..v,.l.n,. 
1„„    ,i„.,v   is   not    \rl    a,i.\     four;'   of   |.,'omTSMo„. 

Section  in  Front  of  the  Anterior  Corpora  Quadrigemina.     W  h. ,,  ik. 

,„,,l„|k,.  |,n„s  a„.l  „„.s,.,i,M.,,l,alo„  a,-,,  inxs.,,,,  as  u.ll  a.  tl,,.  s,.n,al  .-onl. 
,,„.  ,.,„„|i,io„  knoUM  as  .hr,nhn,h  n,,<Ul,  s,„.,.,v,.„..s.  ll,>s  ,s  „,ost 
,„a,k...l  in  „ia.n,uals,  Init  is  also  p,vs,.„,  to  a  ,.<.,-1ai„  .-M-n,  ,t,  ,,>u.-k 
,„,-,,,  ,ni,nals.  as.  f,.f  example,  in  frogs.  1.  ..onsists  of  a  tonu.  oon-htio,, 
of  tho  postural  museulatufo  of  tl.e  lu„ly.  mainly  of  the  extensor  rnt.s- 
,.les-  tl,e  elhows  and  knees  are  exten.i,-.!  and  they  resist  passive  1  exin-. 
the  tail  is  stilT  an.l  st,ai.hf.  the  n,...k  an.l  h..,,!  are  retraete.l.  The  eon- 
,,ition  is  ,n„h,nUtnlly  .In-  to  ov,.,aetivity  of  Ike  ,-ellex  tonie  fnnet.on  o. 

,1„.  spinal  ,1.is.  for  it   ,lisap,.ea,s  ul„-n  the  ,,osle,-ior  s,nnal   roots  a,-e 

,,„      Tl,..  r.th.N,.s  that  ,l,T''i"l  ""  ""•  """■  "'■  ♦'"■  ""'-■"I"""'''     '"'■  "■^""'l'''; 

,|„.  U, .i,.,k  a„,l  ..M.nso,-  thia.s,      a,-e  very  |.n„,o,>ne,.,l  in  sn.-h  an  an, 

,„,,l     a„,l,  on   a, un,    of  ,l,e   l,i.J,er  int,.o,.a,io„    ,,res,.„t.   .elh.xes  ap,.ea.r 

„„;  .„■..  al,s,.„,   in  anin,als  havin^  the  .-erelnospinal  axis  ,-„,   low.r  .lo^^„ 

K.„.  ..xM,„|.l.-,  althonuh  sneh  an  anin.al  ea, t  f.^el.  >  e,  «  hen  a  st,n,nlns 

i.  ,,,pli,.,l  that  in  a  norn,al  a„in,al  uonhl  eanse  ,.ain,  th,.  x<"-al  a,-para1,is 
,„„v  1.,.  .A,Mle,l  so  that  a  sonn.j  or  e,v   of  pain  ,s  p,o,l,n-...l.     'I  he  ,-,.-n,l,l.v 

,1...;  no,   atTeH    th,.  ,...spi,-ato,.y   -nnsMes.     .\,-„.r  sn.-l,   : p..,at:o„,   liou- 

„,,,,    „„,„„1    ,...s,,i,a,io„    is   ,nnel,    nior,.    likely    ,o   l„.    „a„„.,l    „    tl,.. 

s,.,.,ion   Is  in    f..,.,„   ,.f  the  an,..rior  -.n-pora   nua.lri-.M  n,i„a   than   hehnni    „. 

Removal  of  the  Cerebral  Hemispheres.  This  f,„„>sl„  >  ns  vv„h  vvha, 
is  knoun  as  a  .h...e,...hrate  prepa.-ation  tl,at  is,  o,„.  in  uhi.-h  the  an,,,,.-,! 
v..,ains   ..vervthinu'  f.'.m.   the   hasal    gan-.'lia   .lou.nvar.l.      Tl,.'   op,.,-at,oi, 

„,,„lu.-s  a  .■"on-lilinn  whi-h  vai-h's  a- nl,,,-,'  „.  the  hahi,s  of  II,-  an.n.ai 

Thns   in  sn,.l,  /,>f,  as  ,l„.  Hlasn.ohranehs,  uhi..!,  .h'pen.l  for  th.Mr  nnpr-ss.ons 


i.FKi:( 


TS    OF    I.XIM.HIMIA'I'AI 


,i;slONS 


841 


v.TV  larr'-l.v  «>ii  111''  M'li^"'  "f  ^""'ll.  ^\''  '•'"'  ''!"'  »l<''''''-"lirati.)ii  ciiiiscs  tli.>  aiii- 

n,;,i  to  hfcniii .ini.ln.'ly  iniiiKil.il.-.     It  .'an  not  seek  foo.l  !)ooausc  thc 

smsc  or  sMirll,  iipc.i  uliicli  it  nntiiiarily  solely  .IcpfiuN,  has  l.ci'ii  dv- 
stn.Nc.i.  In  a  hony  lisli,  on  111.'  otli.'v  iian.l.  (U'.MTfhraliou  ••auscs  very 
little   .liflVicnce    in    tlic   hcliavior   ..f   tlii'   animal,   pn.vide.l   tlu"   tlialiimi 

and  o|)tic  loW.s  have  I n  left  intaet.     It  eontiniially  swims  about  and  is 

ahle  ti>  dislinLruish  edihle  Irom  noiiedible  material. 

In  the  fro,,  Ww  lesult  depends  very  laru'ely  \\\u,u  whether  the  optic 
thalami  have  been  siiiuiltaneoiisly  removed.  Kveii  when  these  struetures 
have  been  removed  aloni.'  with  the  eerelnnin,  tiie  animal  at  first  appears 
very  little  dilTerent  from  the  normal  fi-oj.'.  It  springs  away  when  touohi'd. 
it  elind)s  ni)  an  inclined  plane,  and  when  thrown  in  water  it  swims.  It 
is,  however,  (|nile  ineapai>le  of  jirodiieinsr  any  sp<mtanei>us  movement, 
iiiid  is  in  short  notiiinv  moie  than  an  e\tremel\  complex  machine,  fo- 
aetiu}.'  always  in  exactly  the  same  way  to  the  same  Uin.l  of  stimulus. 
When  the  optic  thalami  are  also  inta.-l,  spontaneous  movements  arc  said 
to  lie  occasionally  observed.  Such  a  fro''  is  saiil  indeed  to  react  on  the 
ai)proach  of  wint.^r  as  normal  frog's  .hi  bv  preparini.'  itself  for  hiberna- 
tion, and  with  spring,  to  resume  its  .letivity  and  feed  itself  by  catehinsj 
insects. 

In  the  hiril.  in  which  the  operation  of  removini;  the  cerebral  hemi- 
spheres is  a  very  ea.^v  on<-.  the  iMoveiiieiits  'tcr  decereliration  may  be 
cpiite  complicated,  j.art icularly  if  the  ojitic  lobes  are  intact.  Such  a 
bill!  is  more  active  than  usual  .luriii!:  dayliudit,  but  becomes  i)erfectly 
still  in  the  dark.  It  is.  however,  unable  to  distiniruish  friends  from  ene- 
mies, and  il  sho',\s  no  fear. 

As  we  ascend  further  in  the  animal  scale,  the  oiieration  of  deeere- 
bratiou  becomes  very  diflicull.  (loltz,  however,  succeeded  some  y.'ars 
.•i<ro  in  removinji-  iiractically  all  of  the  cei-ebrum  from  a  (/w/  by  pcrfori.i- 
Liifr  the  operation  in  time  statrcs  separated  by  considerable  intervals  of 
time.  '!'he  aninnd  lived  eiirhteen  months  after  the  last  operation,  and 
duriii!.'  this  time  it  beha\ed  exactly  like  an  automatic  machine.  All  its 
rcllcNcs  were  perfectly  normal.  It  could  not  distini;uish  objects,  but  a 
briirht  liirht  cause.l  it  to  close  its  eyes.  Durinir  daytime  it  walked  eou- 
tiiiuously  up  and  down  its  ca'^e,  whereas  at  nii;ht  it  would  sleep  and 
remain  pertectly  .piiet.  When  food  was  i>laee<l  in  the  mouth,  the  dos,' 
would  masticate  ami  swallow  in  ;i  perfectly  nornud  fashion,  and  would 
reject  unpalatable  food.  While  asleep,  a  very  loud  soiiiul  mifrht  awaken  it. 
and  when  a  harmlul  stimulus  was  aiiiilied  to  the  skin,  the  ;!iiimal  would 
snarl  and  u'rowl  and  attempt  to  lijiht  the  .dTeiidin-;  object.  There  were 
absolutely  !!"  siirns  of  i)!eas\ire  (U'  of  recojrnition  of  the  person  that  fed 
it  or  of  fear. 


mfi 


,1  >1 
1  • 


K\'^ 


■llli:   ri  S'l 


I  STUM     N'l  ll^"l  ^    ^YSTIM 


,.,,,|j„st.l,..m,....,an,11lM.  vis....n,lr..Hio,,so1,lu-.n,nu.l1o  -- 

,,.,.,   is,., .sihh..     Altluu..J,  in  ,lH.  ir.dM.,.  .p.^  nn,l  m.wu,   .■-■.,, 1 

;;;,.  ....,Ji,,.„.a..i..  ,.H  .ni,..  ....M„.i ^ --'^ '',••;;;;;;  . 

;,„.,,,    ,■,.,,„.   tl...    -rsults    wl,i.-h   lu.v.   ,,us1    l„...u    ..ons„l..H..l     l.a       n 

::,..,.  ....„„.,.   .„•  ,1 ..-..,„•,.,„.      IfHl.x-s   .l.i..h    in    10   low...   ...u^^ 

.„,,,,,,  .,,,v  a  s,MnMl  or  .  lHm,u-s,rnn,l   .rnH,  al^n  nnolu.   n,,lH.  h^ 

f„nnsa,...r;.l,n,l,...l,  win,.i,  i-i.i.l    i..«n.nh    a-  ,1:.  -ul,   o,   ..M.-n-n.... 

,  ,,,„„,,i,„,     TlH.  n..wly  l.nn,  infant   ,s  al,!.  u,  ,.  r,-„nn   l..w.;r  .nnv- 

..M..an,in.  n..u    .nov. n,s  i.  in u,.aral,ly   „•...,..,■.      ""   "'""•"^  j' 

„,„„.,.,,,;,„    ,.,„,.,    „,,..,s   l.-a.lin.    ,l,n,n.l,    -..■.■lual    n-nrons.    .hn.n.l, 
;  ,i..l w  ,...,„.   ,-..,..,ns..s  nn,>    1,-  l.i.l  ..,...,  as  a  n.nit  oi  0,,... 

l„  eonn..-1ion   wi,l,   tin...  ,.xp..nn,..n1s   .t    .s   n.t..n.snn..  to  uol..     1    t 
i,n™  annuals  i,   ..an   n.aWily   1..   .l..n.nns,,.at,.a   tin,,    ,...n^^^^^^^^^^ 
„,„,„„„  „f  ,1,0  Inulu.r  .m  th.  s,.inal   ....nt.rs   ,s  .„   an   ,nln  .,1,  ,>    natuu  . 

,       ,W,at..n,    ,Mno  ...•  tlK.  Il.xinn   MW  in   tlu.  ,l......nln.a,..    ,r.,,    as 

;U,  l.vth,.Tun.U ,.M,.,.,y,nn,.l,   ,.n.l.n...,lul„.nas„,„uus 

su..h  as  tln,t   pr.,.h...M  l.y  a  ..vystal  ..,■  .onnn-m  salt.   ,s  a,plu..      ,M^^^ 

J,  .,.,,  „„.  .,„,,„,„„  ..x,M...is..s  ,.n  tin.  spinal  .•.■n....s  ,s  an   ,nlnln„„>   ..n.  . 
wh..r..as  that  nf  \W  ...T.iH.lluni  is  an'^nn'malmy, 

— ,,    .    ,„.,,    ,, ,    ,;,,,  ti'ii'   iluit   .;i|'-s  lii-l»vcti  ilii.i.ini; 

l\f  li."l   lilt"  \vi:ik  at"il   -■  1  iiMii  aiM  im    i'" 


if 


m\\   ^\ 


[til 


r!i  1 


CIIAI'TKi;   \i  \ 


(■i;i;i:r.i;.\i.  i,(mai,i/,.\i  i<i\ 


(If  iiiui-li  !jii  liter  inactiriil   iiii|i(>itinirc  than   the  cxixTiiiiriiis  in  wliii-li 
ilii'  ciitiri'   .•(•iiluiiiii    is   rciiiovi'd,   as  .jisiTilu'd    in    tlic   l;isl    clinptor,  Jirt' 


tlidsc    ni    \\ 


tiic  ri'sults  ( 


hii-li   various   parts  nT   it    aic  (|rsli-iiyc( 


1  111'  stiiiiulatcd.     From 


•Iiisiuns   niav   lie  iliawn    rri^ai'ilmu:  liie    ini| 


mi-taiit   suliiivt 


if  (■(/•(/);•.//  liiriili:,ilioii.     '['\iv  I'lVivi-  iii-u.lii I  !■>    rriiKival  or  sliiiiulatioii  (.!' 


ill'iM't'iit  |iaits  (,1"  ihr  rrreliral  rmi. 


\    \AV\     rnllsKl 


iiitcx   liciiiu:  s( 


I    aiiait    I'di-   ll 


ilillol    C.I     llir    llh'tiir    niri 


liody,    iilliiis    I'nr    till'    rrciiitioii    and    intcipii-tat ion    <it'    a 
while  (ithcis,  and  llicsc  li\    I'ar  the  most   rxtcnsix  i\  afc 
ciii'ii'Iaticin    Of   associal  inn    nl'    the    scnsorv    am 


•raM>.  s(jmii'  pai'ts  nl'  thf 
haiiisir.   ol'  IIk' 
iTci'cnt    siinnili. 
iniM'iiK'il   in  till' 


it.'.l 


ni   '_'«in'ra 


d    niulin-    rciitcis.      It    iiiax 
1  that;     '\<    Tlic  nici'iiitral   icirion   »(  tho  ('cit'linmi 


cnntams  thr  cfntcfs  ol   liii;li(i'  tlnm 


-ht. 


lie    asi'i'lK 


liny;  iVoulal 


voliititm  iniiiicdiati'ly  in  t'lunt  df  tin-  lui'i-cnl  ral  sidi-iis  contilins  tlic 
cliicf  Tiiotcir  I'fiitcfs,  a  .Tntcr  hfini;  distiiii^nishalilc  I'nr  cafli  limscular 
LTciiiiiinii'  ol'  the  hody.  ■ :'. .  'I'lic  |>osicciil  lal  i-on\  oliit  ion  has  tn  do  with 
thr  ccnttTs  for  thi>  iiniin'diatc  rfccp.ion  of  sensory  stimuli,  the  so-eallt'd 
sensory  eenters.  (41  A  laruc  area  oeeup\  inir  most  of  the  parietal  lolie 
and  part  of  the  occipital  is  undoutitedly  assoi-iatiniial  in  its  fuilotifUi, 
siiiei'  from  it  no  response  can  he  obtained  liy  siimuiatioii.  eti'.  i  •>  >  l>o- 
hiii'l  tills,  in  the  oeeipital  lohe,  there  i,  a  i-eiiter  having:  to  do  with  tho 
rree|itioli  of  visual  iin))ulses.  it;  In  the  upper  eonvolutioii  of  the  tem- 
p(ii'o-si)h('ii()idal  lolx',  is  a  similar  center  for  hearing'. 

These  centers  have  heen  ilitTcri'Utiated  from  one  another  hy  anatomical. 

ox|ierimental  and  (dinieal  research.     .\t  pr.'sent  \>(   shall  conlii urselvcs 

to  the  rriiitiiiK  iitiil  ris}ilts.  'I'lies,'  are  ohtaincd  hy  aMatioii  and  stimula- 
tion, and  ill  i-onsidcriii-/  tho  results  it  will  he  eonvenient  lo  divide  the 
eenters   into   motor,  sensory,   and    noiiread  i\ c. 


In  I 


\1 


ABLATION  OF  THE  MOTOR  CENTERS 

Iti  iiifirnl  (if  fill  inrli.r  from  the  area  which  controls  the  moveinents  of 
a  deliiiito  [lart  of  the  liody  sa\ .  the  arm  will  he  found  to  produce  an 
immediate  and  pi-ofoiiiiil  miiseular  |iaral,\sis.  Tiie  animal  does  not  use 
tlic  |)ai'al\/ed  extremity    I'or  aii\    purpose  whatsoever,  anil   \  <'t    the  mus- 

si:; 


mw 


iSM    ■-• 


S44 


Tin:   CINTKAl.    M'.RVOrs    SYSTI-.M 


^ilii 


I      •.  i 


I  r 


..l.,s  .1,,  n,.t  UM.Ur.,^.  any  ...,.,■.■  .tn.phv  tlum  <•..,  !>.■  iu-ountod  tor  by 
.lisusr  Tlu.  .-xtrnmlv  .I.hs  not  sutTrr  I'nm.  any  of  tin,  nutritional  .l.s- 
,uvl.an,M.s  vvhi-1.  ^^..  saw  sn,KTv..n..  n,.on  .LMrnrtion  of  the  motor  vvu- 

„,,  i„  tin nl:  an.l  lik.-uis..  lonil  -vll.s  a-tions  ..li.Mt...l  l.y  stnunlatn.n 

of  tlR.  lo.al  .■...•..pto.s  a.v  ,..MlVrtly  n..imal.  A  pinprirk.  for  .nstan.c. 
causes  tlie  usual  llexion  rellex. 

\ft,..-  soUH'   weeks   the    li.nl.    he-ins   t..   reeover   au.l   ean   he   used   m 

volitional  nn.vem.nt.     K v.ry  ,Mpi,lly  ,.rot.MesM.s  until,  n,  the  ease  ot 

,1„.  hi.'her  apes,  it  1 ml's  ahuost  e.m.plete  i.,  a  little  over  four  n.o.iths. 

1,   oeeurs  earlier  in  th-  lower  aninn,ls.     When  a  eenter   is  .lestroyed  on 

the  eer..l.ral  eortex  in  the  ease  of  n.an.  only  partial  r» very  takes  i.laee 

So  that  in  ^-eneral  ue  ,uay  say  that  the  hi.'her  the  animal  in  the  aunnal 
s-ale,  the  less  eon.ple.e  will  he  r.Tovery  fr.nn  the  paralysis  pro.luee.l  hy 
eorehral  ahlatioii, 

R,...„r.lin-  llu  >,„t„n  -/  lln  m.r.ni.  several  possihilities  exist;  either 
,1„.  ,;;.rve  e,.nter>  be..oM>e  re,..M.Mate,l  in  the  destroyed  area,  or  the  eor- 
respondin..'  are;,  of  the  opposite  h,.niispher..  or  >oine  other  l.art  ol  the  same 
.,..i„isph,.re  or  the  hasal  ..'andia  assume  the  funetion.  Kvnh-nee  has  heen 
furnish.d  hv  Sh.  rrin..'ton  and  (iraham  lirown-  ti'iidm-  to  show  that 
,l,e  last  of  these  is  the  most  likely  ■•ause  for  the  reeovery.  Thus,  ,t  uas 
,■..„,„,,  i„  ^vnrkint:  on   th.  arm   rent-rs  on   the  hrain   of  the  ehimpan/ee, 

that   after  eomph-tc   re.-overy   of  th.   pa-alysis  produ 1   hv   removal   o 

the  eenter  on  one  side,  stimulation  of  th.  area  ihat  had  heen  removed 
.,,„sed  no  movem.nts,  indieati,,..-  that  no  reuM.neralion  had  oeeiirre.l    an.l 

,1,,,,    removal   of  the   eorrespon.li,,..^  .-nter   ,.f  tl M>'-'te   hemisphere 

,,„,„,„„,,  ,'.,ll.,„,..l  hv  paralysis  of  the  arm  t..  whieh   it  e.nresponded,  still 

,li,l  „.,t  ..ause  anv  paralysis  of  ih.  limh  ^^hi.■h  had  n vered   from  the 

,„.,.vious  ..p.ratio'n.  To  s.t  whelh..-  so.iie  other  part  of  the  u^ray  eortex 
,„i„l„  i,;ne  MssuMo.l  th.  lost  fun.tion,  the  post.entral  eonvolution  was 
,..,;,oved  two  months  aft.r  the  removal  of  the  arm  eenters,  Althou«l. 
,,  temporarv  weakness  of  hoth  arms  resulted,  the  voluntary  movements 

^v..re  soon  as  u 1  as  hefore.     These  results  are  of  eourse  ..xaetly  wha 

Me  should  ex)...'  from  the  experim.nt  on  the  do-  already  deserd.e.l 
in  whi..h  th.  .enl.ral  .orLx  had  heen  entirely  remove,!,  ami  th.-  ,-on,'lusio„ 
,hat  w.  must  ,lraw  is  that  th.  l.isal  -an-lia  assuni.  the  fnn.-tu.n  ot  th. 
l,ist  ,','r.tiral  ,'ort,'X. 

STIMULATION  OF  THE  MOTOR  CENTERS 

To  iuv..sl,jral,.  th.  .nVets  of  stimulation,  il  is  found  that  the  stimulus  is 
hest  applh-d  hv  th.'  el,",'tri.-al  i.i.'tho,!.  one  point,'d  .-t.'etrode,  ealled  the  stun- 
ulatinir,  heinu'  a|,pli.Ml   to  the  ar..,   un.l..r  invstiiration,  ar..l   the  other. 


;.  .1     ! 


(■|.IU;11HAI,    !,()(  Al.l/.ATION 


845 


(1    the    iiiditTcrciit    clfct lo.lr    and    coiisistiii'.'    "I    a 


It    pliitf,    liomit 


I)liicc(l  ..11  si>iii('  ..lli<  r  luii-1  <>r  tlif  l»nly,  such  iis  tlif  skin  of  the  luifk.    This 
uiii|)oijii'  iiii'tliod  j.'i\i's  iiuK 
clcflrndcs   lire   ciiiiihiycd. 


h  fiiifP  ri'sults  Ihiiii  whfu  ihi'  (inliiiiiry  liipi 


I'.ffiM 


(•   \\{'  di'sci' 


lil'c  tlic  rcsuits  \\ 


hich 


have 


III"   this    iiic 


iifstiDii    aris 


.liicl 


liccii   iihtailicd   hv    tlic   nsi' 
h' 


I     it     lllilV      lie     \\ 


ell     t(i    conslclc'l 


H'li 


(Iv 


iiaiiii'lv.  Iitiw  (III  wc 


tlial  the  ch-rtiic  ciiriciit  is  real 


\-  s 


tiiir 


tiii-r   the    (M'litrr   iircscnl    in    the   iriay   matter   <if   the    .■..itcx,   anr|    lu 


III'  nniiicrous  nerve 


III 


lliat 


stitnte  the  white  ma 


tter  id'  the  hraiii 


;ii( 


aliiii' 


w 


hiell,    lietweeii    the    l\M>    eleel  rodes,    it     Is    |ihlin    sniiie 


(dcetric  cm  rent  iiiiisl  \):\> 
centers  is  as  t'liliows: 


ri 


le  e\  Idem 


i.r    tlie 


(.   tliat    we  ;ile   leallv   st  illllllat  llli.' 


stiiiiiilatiii 


til 


1  1    Tlie   latent 


I'entets  is  llllleli   Inlltrel' 


d   {< 


ilise   I 


I'diluced  I) 


than  that  w  hiidi  fellows  iipoH  di- 


rect  stimiilatii.n  of  the  white  matter,      i  "J  i    I'lider  deep  narcosis,  as  that 


]iroilnce(| 

ma'ier  is  erea 


idihiral  or  morphine. 


the  1 


iTect   ot'  stimnlatioii  of  the  fira.N 


tlv  delaved  and  altered  in  t\ 


I  hen 


stimulation  of  the 


while  matter  uives  the  usual   response 


A  weaker  ci 


urri'lit   siitliees  to 


stimulate    the   tir 
matter. 


itter    than    that    reipiired    for    the    exposed    wir.ti 


li 


1  (in 


ler  to  deiiioiistrate   the  mov 


tl 


le  cert 


)>ral 


irtex,  it    is  ni'i-i 


arv, 


emeiits   which    follow   stimulation   of 
II   he   inferred    fmni    the   pn'- 


as   wi 


ceding'  reinari.s.  that  the  animal  he  not  too  dee|.lv  aiieslheti/ed.  I'lir- 
thermore,  it  is  necessary  to  he  very  careful  in  adjustin-  the  strein.'th 
of  stimulation  employed,  for  the  results  vary  consideral.ly  aecordin-ly, 
Wlieii  the  stimulus  is  of  the  i>nM'er  intensity,  the  movements  arc  located 
ill  some  i.articular  -iroup  of  muscles, --fm-  example,  those  of  the  thuiiih 
or  of  the  hand.-  whereas,  if  the  stimulation  is  stn.iiL'.  the  movemeuts 
spn'a.l  over  much  lar-cr  areas.  .\s  a  result  of  feehle  or  moderate  stimu- 
lation, it  is  found  that  the  muscles  which  move  an-  those  of  the  opposite 
side  of  the  I.ody,  and  that  the  I  oca  li /.at  ion  iv  n,  r  the  liiu'her  the  i.ositioii 
id"  the  animal  in  the  scale  of  development.  The  movements  are  iierfectly 
coonlinate  and  purposeful  in  chara.  ter.  and  recipn.cal  innervation  is 
evident. 

There  is,  however,  a  marked  dilVenMic.'  in  the  reactions  ohtained  hy 
stimulation  of  the  motor  cortex  and  those  ohtained  hy  elicitin;;  spinal 
reflexes.  For  example,  the  movements  pn.ducMl  hy  stimulation  of  the 
c.M-tex  are  much  more  readily  moditie<l  hy  sli-ht  variations  in  the  con- 
dition of  the  animal,  the  Mood  sui.ply.  the  deu-ree  of  narcosis,  etc.,  than 
are  those  elicited  hy  stimulation  of  re.'cpt.u'  neurons.  A  careful  study 
of  this  difTermee  has  Inen  made  in  reeent  years  hy  Hrown  and  Sher- 
rinfiton.''  They  ohserved  the  behavior  of  two  antairoiiistic  imiseles 
aetiiK'  on   the  elhow   when   the   n'speetiv.-   cortii'al   centers   were   stiiuu- 


!«'■ 


w 


il 


.ill 


1 

i 

';i^'  \ 

■'li  ■ 

t 

u  p;t,; 


■m  ■- 

t  I'i ! 

1  !' i' 

K4(i 


nil    (  I  \TU\I.  M  iivors  sY>Ti  M 


|,,,,.l  „n.l  fnuiHl  llwit  llu.  iMtnii  i...n.Mls  ^^rvi■  viy  vnml.l.-,  tlir  mII.t- 
HT.Tts  iiHl.-lliiil.',  MH.l  iiiliiMli-M  inon-  pioiiiiiiont  tl.an  .-xnlMtioi,.  Mop.- 
..vrr  tl...  i„l>iMtion  vvns  ,„..ro  .,r  less  in.lcpon.l.'nt  ..f  tlio  simultiiii.'..iis 
,.x..it„tinn  ..!•  Ilir  ,MntM!.'.M,isti..  nius..l...  in  wlii.-h  ivspcct  it  tli.Mvforo  .1.1- 
tVn..l  iVoii.  tlio  tvi-f  .xliilMt.-.l  il.  tlx'  m-ipioci.!  mwrxiiUou  ..f  tlu-  spin,,, 

,,.,.,,,.     ,,,.,.    ,„ S14».       N".'    "<l''     II'"     f'Mllts    oM;.il..Ml     I'l 1     i.     H.v.Mi 

,,„!ii,,l  ,,,„t..r  m1«mvs  11...  snin,.;  tl.„s,  if  m   p,.int   -.'ivin..'  a   .•.•I'iMin    n- 
.,„,„>.•   uns  ^tii.M.l.-.t.Ml   i,M.n..,li;.t,l.v   .ft..'   p.-viuiis  slim.ilnt ......   11..-   ..; 

suit    «;.s   oft..,.    n.v....s...I:    if   it    XM.s    ii.l.ihiti..,.    i..    n.'--    fi>'s1    n.stai......    .t 

,„i..l„  I x..it,..n,...  i.,.......li=.t..l.v    .ft..ruM...l.     I'..it   if  suni.-i....t  ti.M..  uas 

,,ll.,u...i.  tl...,.  11...  ...sp..,.-  was  MluMvs  .,f  11...  same  U.n.l. 

15  .   ,.„,„,,a,.ii...'  11..-  ..nV.-l    ..f  si...,.11am...us  slimulatio.i   of  a„   atT..,..-.it 
spinal   n.ot   a,..l   of  a   II..Nio..   o,.  ,.M..,.si..i,   point   ....   llu-  <-..t..s.   .t    was 

,.„„n,l   11,;,,    tl,..   sli,„„lalio.,   of  tl...   atr..,...nt    root    wh...i   a    tl..N.oi.    I'on.l 
^.as   iMin.'  sti.M„lat..<l   a,...n,..nt...l    tl...    Il..xi..n.   l.ut    ^^1..■l.    an    .>xt....s.,.„ 

,,.,•„,,   „as  sti.„..lat,..l.  Mi.,.nlar.on  of  tl,.'  nlT..m,t  ..oot  mi'-'l.t  ..Lanu'.-  tl,- 
n.sp..n...  to  ll..xio,..  tl...  ..xa..t   .-..s.ilt  .l.p.M..ri,..„'  ..onsi.l.n-al.ly  .n.  tl..-  r..!- 

.,1iv..  stn....Ml.  of  tl...  tuo  stin..ili.     -n...  --'..„.>,.al  .-..n.-hision  that  may  1... 

'.l,,nvn  f.on,  tl„.s..  .■.•miHs  is  tl.at  tl..  -^V'-An]  f......ti.M.  ..f  tl. ..t.'x  ,s  t.. 

,,.,,,,.s..   ,l„.  .•..nt..,s  of  p.,....ly  spinal    ,...ll..x..s  wl..',.   s,,.-!.   .■.■v..|sal    ,s  .1..- 

.i,,.l,l.^   .>,•   I,... a,.v,     Tl...   .-..t-.x   .l..n.inat,.s   11..'   spinal    r-tl.-x..s.    aiul    n. 

.„,,,„,,,,1   i,   „,:,v   1m.  sai.l  tl.at    its  main   ..flV..t   is  i..l.,l.i1or>    ...   nat.m..^ 
"  It  is  ,.a,ti....'la.h  l.v  tl...  .IS,.  ..f  tl...  ni.th.Ml  of  moderate  electrical  stimu- 
lation tl.at   ..xa.-t   lo,.ali/.atio..   l.as  I,.-.,,   u.^'k,..!  ....t   ....   tl...  ....ivl.ial  .■...•- 

,..x       \s  uo.,1.1  I...  .■xp....t...l.  tl.is  !n..ali/,a1i.m  is  n.n.-l.  I..-  -LIim".!  ami 

,,,.,i„i,.,   •„,    „,..   lo«..,-   as   ..o,Mpan..l    wit!,    tin-   hhjhcv    an....als.      Tn   11... 

,,i„,,.,,.  ,p.,s,  it   l,a>  I.......   f..u„.l  tl.at   tl.."  i.i"t..,.  -•.■i.t-Ts  a,...  l.m.1...1  t..  a 

„,;,.„„    M,.ip  ..f  .-n.t.x    i.a, Iia1..1y    i.._   fn.nt    of  tin-    W..1 W"    l-s..ro 

,1,,.   |,i-. nl.'al   a|...ii.  as  it   is  .-all...!.     '  \'\'-i-  -"* '' 

K,..„n  tl...  a.-.-o,npanyi..^  Ilun..,..  it   .  ill  1.-  not...l  tl.at   11,.;  ......t...s  a,.. 

,,,,,,,....1  f,...n,  I...I0W  ,.pua,-.l.  in  tl,..  r..v..,.s,-  .-.■■l..,.  t..  tl.at  .„  w!,,.-l,    L- 

,„„s..„la..  U.-...IPS  ...•.■u.-  in   tl...  I>...ly;  tl.at   is  to  say.  tl.-   ta.-.  i..-.' <,  |-  <••• 
.,,,  l,.,,„..,l  lou..st  ....  11...  .•...■t..x.  an.l  11...  1--'  lii^'lH-st  np.     It  w.H  furll,.... 

,„.  „„„„,  „,„  ,1,..  ..xt..nt  .,f  11...  ■.•..nt...-s  fo,.  II,.-  „...•!<  an.l  !..»!.,..._  is  v..,-y 

„„„.l,  ,.,...at,.,.  than  f..,-  tl,..  I....ly  ...•  l-'..  H-t    fo.'  tl...  a,-,,,  l,.-..-,'  int...n...- 

,li,1.,      Il    is  not,  th.n.f..n-.  tl,.'  ..xt....t   ..f  tl...  ,.„,s...,lar  Uss.u-   Hiat  -I- 

,.,,,„i„..s   ,1,..    si/.,    of   11,..    .-..i-ti-'al    a,...a    ....ntn.llinu-    its    ,„ov.-i,..-nts.   l.nt 

„„.   tvp..   o,-   -on.pl.xity   ..f   tl...   ,„..v..M.....ts   that    .1,..    ....i-l.-s   p.-..lo.m. 

Th,.  .'. I..X  .,M,v,.:.,..nts  ,.f    Ih..    1..n..',i..    ai.,1    tl...    v.-i.!    -I'-ls    .-vi.U-ntly 

,,..,„i,.,  .M.,.at.r  .-..li.-al  |...pn.s..n1ati"..  tl.a.,  .h-  U..-  -'..a.-s..,.  m..ve..u-i.ts 
^  ,.,,,,  1;,;..,,  ,„ass  ,.f  „„.s..,iiar  tiss,,..  ..f  11,..  tn.nk.  Tho  ....nt.Ts  oxt.M,.l 
.wha,    np..n  thr   ....-Mal  asp.'.-t   ..f  th..  l.,-ain,  In.t   ...■.-ui-y  In-'-e  only   a 


(  I  Kl  HUM.    I.n(  AI.I/.ATloN 


847 


vei-v  sn.Mll  part  ui  tlic  sup-iliriMl  -rM>  nmtt.T.  Tli.y  .•st.iul  ..l.o  into 
tho'tiNsmv  (.!■  IJulniMlo  and  tli.'  ollnr  liNMins,  an.l  tl„.  .'Xtriit  nf  Ih..  ox- 
rital.lr  aiva  wlii.'h  is  tlnis  Imiif.l  away  in  Wn-  lissu  rs  may  .xcMd  that 
nil  till'  t'lci-  surlac-r  111'  till'  iiiMiiispiii'ics. 

If   will  W  iio<.'.l  that   thiiv  aiv  tin,  niihrs   for  th>    movcm,  nt.i  of  the 

,,„,    ii.    thr    rnmtal    l-l..'    isulati'.l    l'i"ia    tin-   inntor   afca.   ami   tho 

..th.T  at  11m.  tip  of  tlir  ipilal   lol,...     Til..  l-nr.n.T  is  the-  in.iti.r  .-..liter 

for  tl.o  ,.ni,iuL'at..,l  iiiuM.ni.iits  „f  tli-  two  .-y.-lmlls,  wl.or.-as  the  latter 
fiiMi.tii.nat.s  ill  assiM-iatioii  xvitli  IIm-  sn-.-all.-.l  visual  .-.-nt.T,  which  rr- 
,...iv,.s  til.,  \isiial  iiMpnssi.Mis  a. Hi  tiaiismils  th.-ni  t..  ..th..r  puits  of  th.> 

Anui  »^V»3ina      Sulcus  centralis 

Abdomtn 
Chest 


Shoulder 


eibo¥t 
Writt 


ves 


Fingers 
« thumbi 


■loiure     ^   ^-^ 
ofJ»w    Op%r\ir 


Vocal cordi 


Sulcus  centralis 


Mistic 


1„-    .himr.l"""--    .■l;"!-'.'^    ,!„■    ,,.i-ai..ii    ..1    Ihi'    i.iii. 


f'.      'II]         ii.:t.r      i-i.tir     1,1      i!;  t-     li'.il!!     "t     tiU'     iiiiinii>'"*«>      -..-.■    -   -.      •  ■■        I  ,. 

KI,!.^c''n;m,Kln''n    ■;,';i;,-   ,.'n      n,.;,.  :;...l    -^ s r,H„a„.   „.,v,.„„  .,.-    ..I    ,h r,.. ,.1,„«   mu. 

.  I(.    Kf""!'^.       I  Aim  r    Sin  ri  iiint'iti.  i 

l„,,i„  t„  |„.  iiitiipivt...!  ami  .■..ni.latnl.  Kx<-itati..n  of  the  .-.'liter  for 
,.yc  ini.v..ni..|its  ill  tl...  frnntal  inl,,..  say.  of  th.'  ridit  si.l.-,  .-ausos  c.m- 
ii,._rat..  .l..viati..ii  of  hotli  .■V..S  to  111,.  ,,ppositi.  si.lc,  that  is,  to  tho  l.-ft: 
■111,1  it  .-ail  n.a.lilv  1.,.  shown  that  this  iii,.v..nu-nt  .if  the  .'v.-halls  is  th.- 
nsiilt  of  n.,-ipr",-al  lhii,i\a1  i,m  ,.f  ih,'  ..\tr:i,>,.|lhi|.  iiiiis.-h.s  (pair,-  SI4-. 
Kvcn  at  th.^  risk  «<  n.p,titi,in  w,.  will  ajraiii  .L'scrih.^  th.-  fnn.laim-ntal 
,.xp,.rii,i..|it  that  ■I,..ii,mst|.at..s  this.  AVh.M.  Ilie  .-yos.  as  in  th.-  ahove 
,.v,>oriiu,.p.t.  iMov..  t,,  th..  l..ft.  i1  m.'aiis  that  tlu^  internal  ivctiis  of  the 
vi.rl,t  ev,.  nn.l  th..  ..xt,.iiial  r...-tus  of  the  left  are  eontraetinjr,  Avherei-.- 
tlu-  externa!   n.-'tus  ,<<  tli..   f.iviner  an.l  the  internal  reetns  of  the  h-,1t,.r 


SIS 


TIIK    t  tlNTKAI.    NIKVOI  s    SYSTKM 


,„•..  l.c.M.iiiiii'-'  nciprornllv  iiihil.itr<l.  th."  ulluT  nmscl.'s  paiti.-ipiil iliu 
t  .  a  sliu'lit  .l.'KiTf.  II'  all  111.-  iHTV.s  lo  111.-  .■xlniM.-ulai-  luus.-Irs  nf  tli- 
viirlit  VM'  aiv  .Mil  cxrrpl  Iho  siMli.  "  lu-'l"  Mipi.lirs  llu-  .•sl.Mual  nHuv 
il  will  \w  r..iiM.l  llial  this  fvi-  I.M.ks  prrm.ui.ntl.v  lowar.l  tlu'  n«lit  Mtl''. 
Ilial  is  an  .-M.-rnal  slial.isiniis  is  |.r.Mlii.M.l.  If  imu  llir  iii.'lit  ••"'t.'X  i- 
^tin,ulat.•.l.  Im.U.  rvs  uill.  as  Maw.  incvr  t..  111.'  iH't.  alllM.ui.'l.  \hr 
ri-lit  rv..  will  IM.I  n.ovr  railli.T  ll.ai.  llu'  ini.Mlr  liii.'.  Its  innv.'in.Mit  :,s 
far  as  this,  iDurvr.  inM>t  .•vi.l.ntly  Im'  dr  ■  to  an  artiv.-  inliil.itinn  ..i'  tli. 
rMl.Tiial  m-tiis  iniis.-lr.  f«v  immk-  of  tlir  ntli.T  iiiiis.-lrs  .-all  ai't  siii.-c  tli- 
ii"r\('s  ai'f  cut. 

TIm-  fxpciiiM.'iit  of  .•oiijii-ati'  .Icviatioii  luiiics  out  aiiotlicr  i>ou'l  n- 
U'Mnliutf  .•.■irl.ral  loralizatioii  nann-lv ,  that  ill.'  n.us.-Irs  wlii.'li  or.linaril.v 
a.-t    al.mi:   with    niiis.'l.'s  on   opp..sil.'   M.Irs   of   tlic   1"h1,\.   air    inn.ivat.'.l 

fn.ni    lioUi   si.i.s   of   III nl.ial   .•..ilex       Tliis   a|.pli.s    not    .mly    to   tli.' 

ni.Acni.Mits  of  til.'  .'V.'s,  lull  t..  111.'  r.'spiratoiv  an. I  ..lli.'r  niov.'iii.nts  of 
11,,'  n.'.'k  an.l  IninU.  |).'>l  ru.'li..ii  ..f  tlic  tnihk  .'.'MtT  "f  tl"'  .'.■r.'l.ral 
.•oit.'X  on  ..11.'  si.l.'  .lo.'s  n..t  pn..lii.'.'  anv  i)anil.vsis,  wliilc  stii.iulalion  m 
this  r.'irioii  i.r.Mlu.Ts  an  ..inal  in. .vein. 'lit  .m  l..>tli  si.l.'s.  We  may  say 
lli.'n'f..!','  that  l.ilat.'riill.v  a.-tiip.'  uiiis.-if>  arc  inn.r\  at.'.l  fr.mi  Imlli  m.I.'s 
of  till'  .'ci'iln;'  .-ortcx. 

The  iiiovcni.  !,i     pr...lu.'.'.l  hy  stronger  stimvdation  ..f  th r.'l.riiin  .1-. 

n..t  niiiain  l.H'aii/c.l.  an.l  ih.'y  p.'isist  f..r  s..nu'  time  after  th.'  stimulus 
hM^  li.'.'ii  r.'m..v.'.l.  Still  furth.'r  incrcMs,.  in  the  str.'iisrth  ..f  th.-  slimulus 
may  .-aus.'  the  .•ontra.-ti.u,  I.,  spr.'a.l  until  it  atT.'cts  all  i>arts  of  th.' 
h...iy.  -ivin-j  ris.'  t..  a  .'..nx  iilsioii.     Thcr.'  ar.'  tw.:  typ.'s  of  .•ontra.'tion 

.luriiiu'  this  coin  ulsi.m.  th.'  tii'st  Lciii'-'  a  str.uiy:  toni.- tia.'li.ui,  -vvhi.'li 

outlasts  111.'  stimulus  for  som.'  lime  an.l  then  -,'ives  way  to  a  scries  ..f 
cloiii.'  c.Hitra.'li.ins,  o.'<'iirrini,'  at   iiil.'ivals  ..f  fnmi  six  1..  t.'U  per  sc.'.ui.l. 

ami  irra.lually  .^'cttinu'  slow.'r  as  th.-  til  .li.'s  away,     Th. nvulsion   al- 

wavs  starts  in  the  muscle  siroup  r.'pr.'scnl.'.I  liy  the  .-..rti-'al  .'.•nl.T  that 
is  i.ein-^'  stimiilat.'.l.  Thus,  if  the  han.l  area  is  the  seat  of  slimulalioii. 
the  .'onviilsions  lic.'in  in  the  musd.'s  of  tli.'  haml;  then  th.-y  sprea.l  t.. 
the  muscles  of  the  forearm  an.l  sh.ml.lcr  .ui  tlu'  sam.'  si.le,  an.l  then  t.. 
the  fac,  the  trunk,  an.l  the  le-,';  an.l  if  the  stimulus  is  stroiiil  •■nousih. 
they  may  si)r.'a.l  to  the  opposite  si.le  an.l  thus  involve  the  whole  ho.iy. 
This  "march"  ..f  th.'  .'onvulsi.ui  .l.'p.-n.ls  up.ui  the  ..vertlow  ..f  th.-  stiniu 
Ills  on  to  the  vari.ms  .'cnt.'rs  of  the  l.raiii,  ami  th.'  i-atliways  ihrniiiih  whi.'li 
it  occurs  seem  to  he  locate.l  in  the  suli.'ortical  .'.'nters,  f.ir  111.'  s))r.'a.l 
is  not  prevent.".!  l.y  isolatins:  th.'  .'ortical  centers  fnun  one  an..th.'r  l>y 
cuts  encir.'liiif;  them,  or  liy  divisi.iu  .it  the  .-orpus  .•allosuiii.  Xrver- 
theless,  the  centers  .lo  in  s.ime  way  h.'cime  involved  in  the  spread,  as 
is   indicated   l.y    the    fad    that    the    complete    excision    ..f   ..ne    ..f   them 


CKHKHKAI.    l.(H  AI.IZATION  M9 

will    «-xclii.lr   tlif   (M.irosiii.ii.liiif.'    nnisculnr   iiri'ii    I'loiii    paitii-ipation    m 
thf  tit. 


CUNICAL  OBSERVATIONS 


'Pile   (^ 


iicifi.ili''    results   nil 


itiiiiicil  li\    cxiHTimi'iitiil  Ntiiniiliitidii  in  iiiiiiiiiiiM, 


a  IT 


ici 


\     silMlllir    to    till'    S.MII|itcillls    (.lls.MM'cl     ill       IIIMII      Wlll'll 


till 


I'l.l'ili 


.■oit.'x  is  stiiiiuiiit.'.i  l.y  the  i)iTs^uf it   "t"  a  in.'MiiiK«'iil  tumor  or  a 

spicuir  of  Im.iic.     Surh  st iiiiulat ion  caiisrs  .•oiitiactioii  in  tlic  c-on't'spoiul- 

ini.'  muscular  area;  tl nliactioii  tlini  sprca.ls  to  iniu'lil-oiim.'  groups 

oi"  miis.'l.s,  anil  may  ultimately  involve  llie  wliole  nnisiMilatuiT  i«l'  tin' 
lioiiy  in  n  eonviilsiv.'  lit,  likf  that  pro.luef.l  in  animals.  This  is  known 
as  ila.'ksoiiian  epilepsy,  ami  it  is  to  I.e  .listim.'uislie.l  from  ordinary 
epilepsy  liy  the  lael  that  the  patient  <loes  not  I.eeome  unconseious  ilur- 
iiiir  the  fit  Like  ordinary  e|>ilepsy,  liowt'ver,  the  Jaeksoiiian  type  is 
usually  preceded  l.y  a  i.iculiar  sensation  of  numhness  or  tiiujlini;  in  llii: 

area  that   is  to  show    the  tirst  < trai-tion.     ( »ne  of  tin-  L'reatest   adiieve- 

nients  of  modern  hrain  sur-rery  is  the  cure  of  .lacksoiiian  fpilfp\^,  I'X 
trephlniiiu'  tlie  skull  over  the  alTected  center  ami  removing  the  m.'liinu'cal 
tumor  or  spicule  of  hone  wliieh  is  iesponsil)le  f:ir  the  stimulation.  To 
enahle  the  sursreoii  to  locate  exactly  the  jHisition  of  the  irritatiiij:  hody. 
it  is  iieces.sary  to  esamine  the  patient  very  closely  as  to  the  muscular 
•rroiii)  wliich  is  initially  alTected  durinir  the  convulsions,  and  then  to 
examine  an  outline  map  of  the  cerclnal  h<Mnis))hcre  iiidicatiiin  the  po- 
sition of  the  various  motor  and  sensory  areas  as  ilediiced  mainly  from 
exp<-rimelits  on  tlie  lii-:her  monkeys  and  verified  hy  the  exix'rience 
"rained  liy  previous  operations.  Toliofraphic  maps  indicatiii-^'  the  sur- 
face iiiarkins.'s  coirespondint;  to  the  various  convolutions  of  the  cerctirum 
must  also  l)c  used.  In  such  operations  the  surt-'eoii  often  has  the  op- 
liortunity  of  experimentally    verifyinjr   the   i)ositi<.ii    of   various   centers. 


■f. 


:sl| 


01 


il ' 


4 


lit 


I  IIAI'TKK   Nt  M 
CKHKItKAl-  I.OC.M.IZATIHN  H'mitMi 

SENSORY  CENTERS 

TliM  tlir  UM.tnr  .Tntrrs  M.v  lo,-;.tr,l  ill  ill.'  iiiriis  Nvl.i.'h  XV  r  Iwivc  just 
,|..srril.f.l  .I.MS  n..t  i.i.li.-.it.'  thnt  th..  '^>■v^<^  ..Us  ..f  tli.'  .•.nt.Ts  n..nnatly 
.loiiiiniit.-  tlM-  tvll.-x  i.i..vn..M.ts  Nv  l.i.-h  tli.ir  stimul.Hi....  .'li-'ils.  Tl..-  n«..l-r 
,,,n1...s,  s1,i.-tlv  s,...akiii..'.  an-  111-  aiit.Ti..r  Ii.tu  .■.■IN  ..f  tli..  spinal  .-..nl; 
„„l  til..  s.,-..mI1..1  iM..t..r  .•.■iil.Ts  ..!•  tl..-  .•.■■•.■I.ral  .-..n.^x  must  n^ally  r.^pr.'- 

M.iit   imtliiiiu'  in......  than   int.Tiiiin.-ial   n.-un,ns  U^^^,^,^u  ll uUnuK  "U'l 

l,.aviii.r  ,„itl,s  iii.^.l   in   nil'N  ni..v.ni.nts.     Tli.^y  an-  ..nly  links  in 

,1„.    loi ..•.•lu-ai    .■liain     u  My.li.,us..s   ..n    tli.^    n-ll.'X    .■.•.•.•l.ral    i.atlnvay. 

\, nlin.'  t..  tliis  ^■H■^^    xv  .■   sl,..ul.l   .•n|..-I    tliat    tlwso   .•.■nt.Ts   nn..u1.1  hv 

„„,  „iiin,at.-  .■.■.•ii.i.^nls  -f  s.i,sati..n.  as  v»  as  llw  .listiilmt..is  ..f  in..tor 
i„,,,uls..s:  s,.hs..rin...t..r.  tli.^y  liax.-  I...1.  .all.^.l  Sn.-li,  Im.nv  .■v.t.  is  tint 
llM-  ..as.-,  f..r  Sli.Trin._'t..n  luis  sIi-.xmi  that  Ili.>  .••■nt.Ts  n...st  .lir.^.-tly  .-nn- 
,.,.rn.-l  in  th..  .•.-.■.■pti.-n  ..f  s..Ms.,ry  in.i.uls.s  an-  i...t  l...-at..|  in  fn.nt  ..I 
111..  I{..lan.li.-  tissiin-  l.ut  inini.-liat.^ly  iH'hin.l  it  In  H"  osr,  ,„ln,u  i,„r,,l<,l 
nr  ,,ostrn,ln,l  r,n,rnlntln„.  Kl.^.-t li.^al  stiinulMti..n  in  this  r.'U'h.n  *]<»■■< 
n.,t  oy,Au'  anv  innii.-.liat.^  .•.■sp..iis...  at  l.^ast  if  the  stimulus  is  not  t..o 
stn.M"      A   ni..v..|iMnt   in-lir-.^tly  .In.'  t..  tl...  .■.■.••■ipt  of  a   s-nsatn-n  may 

1 r„.i,.^.l  1.V  a  stnm..'  sliniulus.  .ii.si  as  is  tho  .-as.,  vvh.-n  th..  visual  .-.■n- 

,.,,.   in    tt,.-   ."..■.-ipital    Inl..^   is   sti 'Jy    stin.ulat.'.I.    lu-.Mln.-in..'   s...-..n.lary 

iiiii\.'in.^nls  ..f  th.^  .'x.s. 

IIist..I...'i.-.  .■N|..-.in..^n1al  mimI  .•lini.-al  .x  i.l.^n-..  has  h.'.'!!  fuinish.^,1  f. 
s„pp„rt  this  l.i.-ati.m  ..f  th.'  --hi-f  s.nv.ry  .-.■nt...-,  Th..  .•lini.'al  rvi.l.m^.^ 
was  furnish...!  l.v  llarv.y  Cushin-^'."  wlm  in.ln.....l  tw..  ).ati..|i1s  in  xvh.un 
this  part  in  th..  hraiii  was  ..xp.,s,..l  1..  alh.w  hii.i  t..  stiniulat.^  it  while  1h..y 

,y,y,  in  ,. ns..i..us  slat...     As  th.^  .■.■suit   ..f  th.-  stin.ulati.m  of  tho  p..st- 

....ntrai  .■..iiv..luti.Mi  .h'linit.^  s..n...ry  iiiipr..s>io„s  w-r..  ..xp.Ti.^n 1.  .-onsist 

i„.„r  „,•  ;,  Musati..!!  ..r  nu„ilm..ss  ..,•  ,l..a.ln..-  t..  ta..tual  in.pr..sHi.u.s.  Lut 

11.,' .iius.-ular    -n.ui.s    .in.l.T«  .'lit    iii..\  ..in.  nt    uiil..ss    111.,    pr ntral    .-n- 

v..lu1i..n   was  stiniulat...l.      Durin-  th..s,.  .novfUKUils,  n.(M^....viM-,   11..   -.,1- 

sations  u.-i-..  ,.xi..Tii^n.-..l  '>>  ll--  l'.tt»'"t  '  ^'-'-I't  t'">^f  ^'^'i'-l'  ■' '"••?••'■'•'"■'■ 

th..  .■hai...'.^  in  th..  p..siti..n  ..f  th..  i-aH  that  was  ni..v,..l.     Tho  sonsati.-n^ 
uhi.-h  an-  thus  n.pr.^s..nt...i  ..„  tlir  r.y\rs  aro  llms..  -f  t..u..h  ,1,s,tui,ih;i 


t  1  HI  HUM,    H'l   M  l/.\llii\ 


8r>i 


i;..li   jiikI   tliM>r    ninlint?   to   tin-    poNitioii    ati.l    lii.i\       ..•Mt>   mI'   llir    liiM-fl.s 

I'liin  nii.l  Miip.  iiitmv  Miisiiti-HH  tin  iM.t   -..  .iii  U,  li;i\ rticnl   r.pr.x'i, 

tiilion. 

Thl'lr    In  >'f  .•mils,    11   rluM.  ilNsncillti.lM   ImIw.mii   •sfiisoiy   iiikI    IIMiI"'    .'.Ml 

(.IS,  lis  is  ilhistri.li.l  in  til.'  .  Np.Tiiii.'iit  .l.s.TilM.l  ds.'W  li.r.'  im.l.i'  tli-' 
li.Mil  ,.r  ii|Ms||i,si,i  ipiiL'.'  *«:'.'^>.  ill  "lii.'li  il  will  III'  iviiii'iiil.,T.'.l  that  111.' 
..,,.,11.1.1. ■  s.Mlii.ii    ..!■  nil  111.'  |...st.'i'i..r  i.H.ls  ..f  nil  .'Xti'.'iMity  r.'mlois  11,.' 

,,Mii  lis  .n'.M'tiN.lv    i,niiil>/.'.l   f..i'  x..lili"iiiil   i""^«' 'I'  ""*  '•   """I''  ''"^'" 

|„.,.|i  lu.l  tli.  iH..t..r  I....IS  lli.iHs.K..s  l„rii  .'lit  AlT.r.nl  iM.piilH..s  an- 
tli..r.-r..i'i'  ii..'.'ssjir>  r.,r  lli.'  .  ni.-i.nl  Mii;ti..niil  .'..liti.,1  ..t'  tli.-  imis.'ilIiU' 
llinv  fllllMlts 


SENSE  CENTERS 


All.lllpts   In    |,>C)lti'   .'Nil.'ll.N     III.'    P"-<it 


Mill    nil 


III, I'.lillll    .Mirt.'X    V^  ll.'l'" 


illlprcHsinli-^    nl" 
.1 


111.-   pi,..ii.'i.iit    sfiisiiliniis     visinii,    li.iiriiiir.   etc.-    arc 


ivc.l  an'  nt   .-.iiii'^''   iimr.'  "i' 


•\piTiiii.iil'<  li;i\i'  to  Ih    perl,, nil.. I  ..ii  .luiiih  annuals 


ijini.'illt    l.i.'iinsc   nl'  tilt'   I'a.t   lliat    lli.' 

NcVfltllfl.ss   snill.' 


iiil'.iriiiali..ii  i-aii  I..'  '.'i.'.iii 


,1    \  annus   pal 


,1  I'l-.iiii  till'  nsiills  n|'  a 


Mali. Ill  iiii.l  stiniiilali.in 
1.  Uni; 


Is   ..I'  III,'   .•nil. A.   al.iatini.    .•aUsiinr.    fnr   .'\aiiip!.'.   .l.'luih. 
,vi, I, .,„•,.    .itli.T    ..1    MiiHln.'ss    ni-    ..)■   .l.arn.'ss,    ini,l    sliiiinlalinii    .•aiisiii'.' 


•Ills  nt    tin-   .'\.'s   nr   .'Ill's   siliillii 


ihir  tn  I 


linsi'  nnliiiai'ily  nl.s.'i'v.'.l   wli.'ii 
III. 'SI'  nivalis  arc   si iiinilalc.l   ill   ill.'   Usual    way. 

Til.'  ,ni<l,l„r„  ,<,il,r  is  l,,.'at..l  in  Hi.'  l.a.-k  part  nf  Hi.'  sup.'rinr  (.•inpi.i'al 

iv.,liili,in      Sliiiiulalinii  <.f  llii>  ar.'a   in  .•minials  .•aiiscs  a   ).i  icUiii._'  up 

,,r  II,,.  ,..,  ..11  the  nppnsil,'  si.l,.  as  if  111.'  animal  Inanl  a  s..uii.l.  Cliiii.'al 
iil,s,.|'\  ali.iii    lias    .•.iiiliiiii.'.l    lliis    .•..n.^lusinn. 

TIh'  rlsioil  ,,i't<r  is  ]i,rn\<;]  ill  Hi il'Hal  1..1..'.     It  is  inii...i'taiil  I.,  n- 

p.at  auain  Hial  tli.-n'  an'  I".,  .-.'iit.'rs  ,,n  lli..  ,...i'..l.ial  .'nit.'x  .•..n.'.Mii.'.l  in 
Nisi,,i,:  111.'  fr,.nhil  r;,„.il  r,„hr.  In.'al.'.l  -  u.'  luiv.'  s..,.n  in  Hi.-  fn.nfal 
!,,!„..  an.l  Hu'  s,,,^all..l  rlsa.il  ,  , -./.r  ils.lf.  V»',x\»\  in  Hi.'  ...•-•ipilal  Inl"'. 
Stiiinilatinii  ..r  111.'  in.nlal  visual  .•.'iil.r  i.n,.lu.-.'s  a  i.n.mi't.'r  niovomont 
,,r  H„.  ..ys  tliaii  .1...S  sliiaulalinii  ni'  Hi.'  ...•.•i|>ita!  .•.'IiI.t    ni.-i.-atin-r  that 

,]„.  iVnii'ta!  .•.•nt.T  lias  tl,.-  i.MiiM.liat H'..!  "I"  Hi.'  nins..ular  ni..\  oni.'iits, 

wli.nas  Hi..  ...•.•ipital   Inl..'  is   pmliaMv    .' .r 1   in   tli.'   a.ljuslin.'nt   ..f 

,|„.  iiHiM.iilai'  n'a.-ti.,ns  u!,i,-li  an    ,„■,■.  ssaiy  in  ..niitn-llinir  Hip  <'yp  ninvc 

„ts.    s.,    tliat    Hi.'    ..I'j.'.'ls    ina>     I..'    i.n.p.'ily    xi.'"'.l    Mini    .i.i,l._'ni.'iits 

,',,n,i,<l.  l.y   Hi.'  .'Nt.nt   ..f  Hu'   ni..v,.|M,.nls.   ..f  its  .listan.'.'.   p.isitinn.   ct.-. 

.,-!,,..   .,,,,„,,]   ,.<-si>nnso  1(1  stimu1ati..n   ..f  tli<> il.ital   .•.'iit.'is  sln.ws  Ihat 

,li,.  1„1„.  n„  nil-  si.],,  is  .•..iin.'.'t.'.l   «iHi   111. nvspnn.lin.,'  Iialf  ..f  oa.-li 

...fi,,;,.   tl„.   f,.,v,'a   .'.'iilralis   l...iii-    linu.'v.'i'.   .•niiii.'.-t.'.l    with    l.-Hi    l..l"s. 


:5' 


il 


\\ 


IIMIl 


li  ^i^*i 

.      ,1  .    ; 

•    ^i  :  :' 

il:;.  ^' 


S.VJ 


in:  (iNruvi,  M.innrs  svsti;m 


ASSOCIATION  AREAS 


Tlif  lM-illi,-int  .Milr.,.iM'  ..r  111-  n-    !vlic>  n.n.lu.'t.',!  hy  tlu'  ...MMTimciitiil 
,i„.lli.,.|   ill  .■iiMl.lin-  us  1..  lorMlc  111.'  .-liiH'  n...t.ir  iiml  sonsm-y  amis  of 


M T  l.i;  ..M.i 


Fll!.     -'-'" 


/ 


\  ,-»T 


»♦/ 


'i:  '^k 


\  „■;,:;->, hi. 


^1,     ,ii:)    r.  ■■!     I  ,iiN        1     ll'.      ..  •.  l.i.il     ■"rli 
,,iit    111,       '  ;;.  .I'.iv  n    Ir-K      M..ii  "U   I 


I'.ir     .I<~.ripli"" 


,h.'  .•■•iv!,!-!  .-.■t..x  l-iiv-  yrt  nii.-li;iil.Ml  iliosr  vMst  iir.'as  lyiiiir  hflw.'-n 
1li..|M  wliirli  -lo  not  n.s|.,.ii.l  iti  ,iny  .iHinitc  xviiy  lu  Mrliti.-inl  si  i,nuliiti..n, 
ati.l  the  iililjilioii  <,(  wliii-li  n-sults  only  in  iiiinv  or  l.'ss  iinli'tiiiil.'  synip 


(■i;ui;ni!Ai.  r.diAi.iZATiox 


853 


tdins.  Ill  uidcr  to  (liM'c.v.r  tlif  '  x.ict  riiin-liini  "I"  tlifsf  jiiki^.  il  lias  '.lei'ii 
iio.-essary  Id  .•inploy  an  cntiioly  din-.Tciit  iii..tln"U -tliat  <.f  liistolof?!.' 
1111(1  ..iiil.rynl,.u'i.-  .•xaiiiiiialion.  Wlu'ii  111.'  i.afl.Tiis  ..f  tho  may  cnrtrx 
HIV  coiiiiKiml  with  tlir  lialiits  »f  tlic  animals,  in  <lit1Viciit  fin.ups  of 
jiiiiinjils   iphylojiciHti.-   Mii.ly.   or   .■v.'ii    in    dilV.T.nt    iiidivi.luals   of  tlii' 


.Mf  B-f  D  *Tt    '■■R!tt.S"'*V 


■    I :■..!.    ..i    11..     I  ll"  I     !'!■■'•■'     ■>"■      ->■■  -'^ 

Iran     lil.rii.     .l-     .|i  l.  :  Iini...!     liv     111..     li,l.  r. 
i.l.r.iM.:;  ...  Iv      ,..l:i.l'.-.     I'll     ll">      i.fh..ll. 
l':,,;,,!         ,|.-,.n,     M.„l,>u.     ..1...     >■ .1.11    ■ 


..!,  lllC  ..Ut.T  I  .1  '  :itt.l  llU'^i.ll  I  /.' ' 
-tf.ulin.  "I  ll'.l-  l.ill.N.  ri"'->' 
Uiiiir:.!     «:iv     l.-w     III.'     ."it.v     i> 


si'fnl     kno\vliMl'_'i'    riinci'VMint: 


siiiiii'    ;rroiip     loiiIoL'i'ncti.'    ^Indv    .    nim-li     u 

,,.,p,l.r;il    lu.'i.ii/ation    .-an    aUo   li..   ..Min.'.i.      In    tlu'    Imman    animal    much 

proirivss  is  hoiin:  nia.l.'  hy  .■omi)atinn  tlic  stnictmal  pattoni  ..f  thi'  cor- 


|i!l 


■  •!! 


i     U\ 

i 


fr 

i"fr 

Hr)4 


iiii;  crNTim    \i  uvms  :^vs'ri:M 


;■  !  I 


i 


fi.  li 


.  !, 


H 


i-n  I, 

I :  I  ^ ! 

rl' 

'till 


hi  ^'  I 


1l .  ■  i 


I,  ! 


t.-\  ill  (liflViTiit  pails  .il'  the  iiciriiial  lirain  willi  tliat  I'oiiml  in  tlic  l>iaiii 
c.f  psyclidiiMtliii-  iiKJiviiluals  whose  iiu'iital  s\  iiiptoiiis  liave  liecii  i-an- 
tiiUy  stiulicil  Kft'orc  (icatli* 

r.ir   till'   pinitipsi-   of  tliis   work    it    is   iiociss;n-y    to    iccoL'iii/.o   scvciiil 
lamina'  or  laxfis  ..t'  ihtvc  <-<'I1s  and  now.'  liln'is  .'oniposiii!,'  tlic  coiIcn 
Till'  most  piartical  division  i>  rcpiTscntp.l  in  l-'iir.  "-JU,  ami  is  as  follow  > 
ilia  iiaiTow  snpciticial  laxfr  of  nerve  (il)ers,  with  a  lew   scattered  ee!U 
tlie  outer  fiher  lamina  or  moleenhir  iaver  ;  r2>  a  iiiiK'h  wider  layer  of  small 
medium  and  larye  pyramidal  cells     the  outer  or  iiyramidal  cell  lamina, 
i:!'    a   layer  ..f   •iianides     the   middle   cell    lamina:    i4i    an    inner   layei 
of  nerve  tihers.  sometimes  eoiitaininu'  laru'e  solitary   cells   i  I'.etz  eells^ 
the  iniu'r  liher  lamina:  i .".  •   a  layer  of  ])olymorphic  cells     the  inner  cell 
lamina.     This  live-layer  tyi)e  nnder<.'oes  structural   moditicalions  in   the 
dilTerent    re;;ions  of  the   coit<'\,   and   these   modili<-alioiis    possess   a    di- 
liiiet   functional  siiriiilicance.     The  onl\    part  of  the  brain  in  which  tlnv 
can  not  lie  rccouiii/eil  is  thi'  hippocampus  and  th.'  pyriform  lolie.     P.ased 
on   the    relative    thickness  of   tliese   layers   maps   of   the   <'erehral    cortex 
have  lieeii   piciduced.     The   iin-st    impoitaut   are  those  id'   Urodmaun   and 
Camithell,    (if    whi-h    the    lattei-    is    reproduced    in    Fiji'.    'J'^l.      Two    le 
yioiis   can    he    \cry    il.'linitely    located;    namely,    the    preiMiitral    oV   Bet/ 
cell  area,  and  the  visual  or  visuosensory  area  of  ( 'aiiipiiell.     Sui'l-ouudiliL' 
the    visnoscnsorv    area    is    a    detinite    \  isuopsx  idiie    area,    and    similarl\ . 
hoiderini;-   on    tlu'    preeeiitial    is    the    so-.'alled    intermediate    i>ivcentral 
area.     At  the  \  ei\    front  of  the  frontal  lolie  is  the  iirefroiital  area.     I'osl - 
rentral    and    intermediate    postcentral    ar.'as   are    indii-ated.   hut    the    re- 
mainder of  till'  center  is  undetiiied. 

K'easonini:  frcuii  iihylo^'enetio  and  ontovenetie  data,  we  can  assifin  to 
ea(di  of  these  lamina'  a   functional  si^'uilicanec.  which  aeeordinir  1i>  Boi 
ton  is  as  follows:     The  outer  cell  lamina  :|)yramidal  cell  lamina)   prolia 
My  eonstitnt.'s  the  ••hi<:hcr  level"  l.asis  for  the  carrying;  on  of  tlie  liiudier 
or  psychic  eer.'hral   functions.      It   is  a  prominent   feature  of  the  human 
cortex,   the    last    cell    layer   to   hi'   evolved,   and    the   one   which    underfTois 
retro;.'ression   most    readil.v.      During'  di'velopment    it    rapidly   attains  ma 
tnritv  in  the  visuosensory  area,  next   in   the  visiiopsychic.  and  only  later 
in    the   prefrontal    rc/ioii.      In    the   \  isuopsychic    area    it    is   i)ractieally   ol 
the  same  depth  as  in  the  visuosensory.  whereas  in  the  prefrontal  i-eu'lon  it 
.levelops  aeconliiij.'  to  the  menial  capacity  of  the  animal.     In  patients  e\ 
hiliitin^'  syni|)toms  of  ileinentia   this  la.\ei'  of  cells  is  distinctly  di'lieient 
Those   facts   indicate  thai    the  onti'r  or   jiyramidal   I'cll   lamina   "suliserv.'- 
the  jKsychii'  or  associational  fnuctions  of  the  cei'i'lirum"      (Bolton). 

•  \t,  .s>.I.M,l  ;,>>,. util  ..f  Hm-  i.luM.il.iKH  li.i-i'<  f.ir  -uli  «-ik  is  liivfi,  l.y  Ilolt.ni  in  l.."iiar.|  11.11  - 
I  iiuh.r  .-hh. ,«.,-<  M.  |■l,^„  l,;n.  Wo  b.ivr  ..klI.'  (r.r  i,~.-  -f  lliis  .irlMie  in  Uir  |.rcs.-t.t  r.-vi.w  .v  .1 
«...ii.i    Mrn.iRlv    r..nmni.n.|    iM    |..  ui'.il     l.v    .inv    wli.i    iii.iy    .l.-iie    fiirlli.  r    iiiformati.m." 


n;Ki:itu.M,  i.cmai.i/.atiox 


85:. 


Tlif  ui'uMUi  cfll  liiiuiiia  is  miich  liy|)('rtn.|)liif(l  in  the  Mi-calicd  projcf- 
tioii  areas  of  tin'  .•.•rcliniiii  fur  cxainpli'.  in  tlic  visuoscnsory  area  (s.'C 
Fig.  20),  wlicie  it  is  sc.  tliicli  that  it  is  tisiialiy  (Icscriiied  as  ix'iii','  (livided 
into  two  parts  l>y  a  narrow  filifr  band  (ilic  lino  «(  (icnnari).  Diminution 
in  tli(>  layer  oci-nrs  in  the  visnosciisory  area  in  lontrstandinR  cases  of 
atropliy.  "It  seems  tlierefore  primarily  to  snliserve  tiie  function  of  re- 
ceiviii','  afferent  impressions  wlictlier  these  arrive  directly  from  the  lower 
sensory  neurons  or  indirectly  tlirous.'li  other  reirions  of  the  cere])rum." 

The  fifth  or  inner  cell  lamina  is  the  first  to  tiocimie  differentiated,  an.l 
it  is  of  extremely  constant  depth  in  the  adult.  It  is  not  iiuich  alTected  in 
amentia,  unless  when  this  is  very  severe,  as  in  patients  who  are  unahle 
to  carry  on  the  ordinary  animal  fum-tions.  In  short,  "it  suhscrves  the  lower 
voluntary  and  instinctive  activities  of  the  animal  economy"— (Bolton). 
Takiiifr  the  rcMdts  as  a  whole,  it  appears  that  the  rc^'ion  of  the  eortcx 

hohind   the    K'olan.lic    lissof nsists   of   sensory    areas   and    association 

areas  wliidi  may  he  immediately  coi cted  with  them  ( visuoi)sycliic  and 

internu'diate  postcentral  i  or  more  removed  (in  ));irietal  lolie^.  The  por- 
tion in  front  «{  the  Wolandii-  lissure,  on  the  other  hand,  contains  the 
elTei'cnt  ai'cas,  of  which  the  precentral  may  he  rcjiar'-d  as  of  lowest 
L'radc.  The  motor  dischar<res  from  it  are  conditioned  upon  impidses 
connnfi  partly  from  the  adjacent  intermediate  pr>'ccntral  area,  in  whidi 
auain  are  clalioratcd  those  received  from  the  sensory  areas,  and  partly 
from  tho.se  eominir  from  the  ]>retrontaI  reu'ion,  which  is  the  most  com- 
plex zone  of  assoeiation.  This  last  is  indeed  the  supreme  dominatinu' 
area.  It  coordinates  or  intejrrates  the  activities  of  the  other  association 
areas  and  nmy  he  considered  as  the  seat  of  the  intellect.  The  evidence 
for  this  hi^li  evolu1ii>n  of  the  i)rcfroiital  area  is  very  stronsr.  It  is  the 
last  )>or1ion  of  the  cortex  to  he  evolved  and  the  first  to  undergo  retro- 
•.Tcssion.  In  idiots  and  imheciles  the  thickness  of  the  pyramidal  cell 
layer  in  this  rejjiion  is  directly  i)ro])ortional  to  the  mental  jtower,  and 
in  dementia  desrrces  of  retro-rression  occur  that  vary  direotly  with  the 
existing  s.'rad<'  of  dementia.  In  normal  hrains  this  layer  is  the  very 
one  which  varies  in  depth  in  dilTercnt  individuals,  Alon-r  with  its  hiiih 
develoitment  in  the  hrain  of  man.  as  compareil  with  that  of  other  ani- 
mals, fToes  hand  in  lian<l  a  izreat  increase  in  the  other  association  areas. 
ThoU'/ht  is  the  i>rodMct  of  inteirration  lietwcen  these  various  associa- 
tion areas,  and  articulate  and  written  laniruaL'c  the  outward  manifesta- 
tion of  the  process.  It  is  owiufr  to  the  relatively  <;real  extent  and  com- 
plexity and  constant  development  of  the  prefrontal  lol)e  that  man  so  far 
excels  even  the  lushest  apes  in  his  intellectual  activity,  and  it  is  owinjr 
nctii>ti;d  ih'velopniei.t  oi'  this  l"he  that   individuals  dif- 


f(.  the  reh 


!iv< 


fu 


Ill  ■ 


fei-  from  one  anothei'  ni  tlu'ir  me 


ital  p<i 


iiiH^  I  '''I 


1  (., 

t- 

nil'  i 

■iWi 


m  it' 


i     ',    !l 


CIIAPTKU  XCTH 

(■(.NDITIONKI)   AND   INTONDITIONKI)   RKFLKXKS 

In  slu.lvinu'  llu"  .■.■iK-x.'s  in  tho  spini.l  animal,  «»•  Lave  seon  that   tlu- 
otlVct   of  a    civn.   sti.nnius  or  of  .liflVnM.t    stininli    is   pir.lu-tal.lo   u.tl. 
al.solute   .•iTtai.itv.     Tlu'ir   is  a    fatality   in   tlu-   irspouses.     When   the 
hi-her  .-enters  are  inelmle.l  in  the  velh>x  are.  the  reth>xes  heeome  modi- 
lied  or  inhii.ite<l  l.y  events  oeeuirin-  in  other  parts  of  the  eentral  ner- 
vous svsten.  and  the  resr.lts  eon.e  to  he  more  and  more  unpredietab  e. 
Th..  rellex  e.m.es  to  he  a  eonditioned  retU-x  (Pavh.v).     Studies  ot   ti.e 
.•innnnstanees   atTeetins;    these   .-onditione*!    reflexes   really   .'onstitute    a 
studv  ..f  the  fun.-tion  ,.f  the  higher  eenters  in  the  hra.n.     Snu-e  sueh 
experiments  n.ust  he  performed  on  the  lower  animals,  we  are  limited  in 
the  invest iu-at ion  to  motor  responses.  f..r  Ave  have  no  way  whatever  ot 
studvin-  the  suhjeetive  sensath.ns  ,,roduee.l.     Tin-  nn.tor  phenomena  l.y 
uhi.h   the  animal  may  eM.ress  its  sensatio.is  ...11   he   interi.reted   hy  us 
onlv  in  tern.s  of  psyeholosrie  ideas  that   in  lar^-e  part   are  der.ved  from 
„„;   ..WM   experienees.      Ohviousiy    the   eonelusions   that    ean    he   .Irawi. 
nre  suhjeet  to  verv  iireat  sourees  of  error,  and  it  must  he  eons.dered  as 
„ne   of 'the   Kn.itest    advanees  of  m.«lern    physiology   that    Pavlov   and 
others  sh..nld  have  sueeee.le.l  in  ev.,lvini:  methods  hy  whieh  we  may  :ir- 
nve  at   eonelusions  reu-ardin-  the  nature  of  eertain  of  the   nile-rations 
„,„   „,,„,  in  ,he  higher  eenters  of  the  nervous  system   preeedn,..'  the 
,„„,„,  nnu.ifestations  l.y  whi.-h   the  animal  exi.resses  .Is  sensations. 

The  methods  en.i.lov...!  for  the  study  of  th.-se  hi-her  integrations  of 
,|„.  eentral  nerv<.us  system  all  dei.end  on  the  reaetions  of  the  animal 
,l,at  are  asso..iate.l  with  the  taUinj:  of  fo<..l.  When  the  food  is  aetu- 
MJlv  plaee.l  in  the  mouth,  it  exeites  a  s.'eretion  of  saliva,  whatever  the 
..in-uiiistanees  mav  he.  This  is  an  uneon.liti-.ned  rell..x.  Suppose,  how- 
,.ver  that  everv  time  fo..d  is  irivn  a  paitieular  s..und  is  made;  after 
some  time  it  will  he  found  that  the  ..eeurrenee  of  the  sound  alone  is 
suni.-ient  to  eause  a  seereti..n  of  saliva.  In  other  w..rds,  a  eond.tione. 
n.llex  has  h,-en  formed.  Similarly,  si-^'ht  or  smell  or  any  other  type  of 
sensation  n.av  he  made  the  exeitant  f..r  the  e.M.(liti..ned  reflex.  The 
seereti..n  now  becomes  psyehie  insteiul  of  merely  physioloirie.  To  .fuote 
H'lyliss-  "Anv  phenomenon  of  the  outer  w.rld  for  whieh  the  animal  m 
,,uestioii  p..ssesses  appr..priate  reeeptors  ean  he  drawn  into  temporary 

830 


COMHTIOVr.P    AND    rN((>NM)l'riONi:l>    KKFI.rXKS 


857 


association  with  salivafv  s.ciction,  so  tliat  it  iicfoin.'s  an  cxfitcr  of  se- 
cretion if  only  it  lias  hccn  lrc<inontly  i)vosont('<l  at  tlic  sanio  time  with 
the  unconilitioiicil  letlex  stiniulns,  food  in  tlio  mouth." 

Work  alonsi  lines  similar  to  that  devised  hy  Tavlov  has  nn)re  recently 
heen  nmlertak.n  hy  students  of  animal  liehavior.  who  have  utilized  the 
aeciuired  hahits  of  an  animal  in  searehini:  for  its  food  in  order  to  study  the 
intlueiice  of  eon-litioniui.'  circumstances  on  its  jirocedure.  The  advantage 
of  this  method  depends  mainly  on  the  fact  that  it  eati  he  aiiplied  to  all 
uroups  of  animals.     In  carryinir  out  such  an  observation,  the  aiumal  is 

,,li,ced  ill  oil >m!.i;rtmeiit  of  a  ea!:e.  fnm.  which  it  is  then  released  to 

a  second  comiiaMment.  the  end  of  which  is  divided  into  two  passage- 
ways, one  IcMJiia'  to  food,  the  other  leadinir  to  some  compartment  in 
which  the  animal  is  |.niiished  f.ir  its  mistake  as  hy  reeeivinff  an  electric 
shock.  Ohjects  such  as  cohoed  lidits  are  ]>lacedin  the  different  pas- 
sair.'ways,  and  the  animal  hy  rei)eated  trial  eomes  ultimately  to  learn 
which  vartieiilar  colored  lijiht  siirnities  the  i)assa-:e  alon<r  which  he 
will  re.  .ive  food.  A  retlex  has  therefore  become  established  conditioned 
on  the   particulai-  colored  li'.dit. 

On  account  of  the  unavailability  of  his  publications,  it  is  impos.sible 
at  ))ri'sent  to  jrive  any  i  oniplete  account  of  Pavlov's  discoveries.  A  few- 
facts,  however,  are  of  such  importance  that  it  is  necessary  for  us  to 
state  them  hen-  as  far  as  we  know  them,  i  S.c  Hayliss.  I'hiiniahimt.)  Two 
mechanisnis  seem  to  be  coiiceriied  in  the  conditioned  reilexes:  (1)  that 
uf  tem;.orary  association,  and  (12)  that  o''  analysis.  Temi)orary  assoeia- 
tio>:  is  well  illustrated  in  the  above  experiment  in  which  the  secretion  of 
saliva  is  induced  by  a  sound.  Temporary  association  of  the  soinid  with 
the  secretion  of  the  saliva  may  readily  be  inhibited  by  all  kinds  of  ex- 
ternal idienomena:  thus,  if  the  do<.''s  attention  beecunes  diverted  while 
the  conditioned  retlex  is  beinv  stimulated,  the  r-sponse  does  not  occur. 
Ill  a  dojr  that  had  been  trained  to  screte  saliva  to  sound  of  a  par- 
ticular metronome   beat,    inhibition   urred   one  da.     because,  .just   as 

the  do!,'  was  beiiii:  i)resente<l  with  the  food,  the  laboratory  servant  made 
a  noise  (uitsid.-  of  the  buildiii!.'  which  diverted  the  animars  attention. 
The  eoiiditiniied  letlex  m;iy  also  be  iiiteifi're<l  with  by  internal  inhibi- 
tion, which  is  illustrated  by  experiments  in  which,  after  a  do;:  has  been 
trained  to  respond  to  a  jjiven  .■onditional  retlex.  M'veral  occasions  follow 
when  food  is  not  jriven  to  the  animal  after  the  iiarticnlar  sensation  to  which 
it  has  been  trained  to  re-pond.  The  condition —fm-  example,  a  sound— loses 
its  efi'eet.  This  is  internal  inhibition,  but  it  is  a  temporary  condition 
since  tlu'  rellex  returns  of  itself  after  a  iieri.nl  of  rest. 


These  exp( 


riments  illustrate  what  is  mean 


t  bv  the  formation  of  tem- 


j 


porar.v  associat  ion- 


rcurrinj/  in  conditioned  reth 


'xes,  but   in  order  that 


Hr.8 


•nil.    (KNTKAI.    N'KKVurs    sYsilM 


ii'.      M 


d'     !■ 


;;    '^ 


It  ■•! 


Mi  i ; 


tluTO  inav  1.0  a  fine  discriiMinatioi.   l.Hw,..u   tliosr   stimuli   wliu-h  slmll 
aiul  tluist'  which  shall  not  sorvo  t..  call  |-..rth  the  c.Hulitioucd  voflcx,  an- 
other mechanism  becomes  involve.!     that  of  anal.i.^is.    This  is  perfornuvl 
l,y  a  sense  oifian  the  function  of  xvhich  is  to  separate  an.l  .listin-uish 
the  complicat.-,!  phenomena  of  the  outer  worhl.     For  example,   it  has 
hcou  proved  that  small  dinV.cnees  in  the  pitch  of  a  musical  note  mav 
determine  whi'thor  or  not   a   eoM.litioned   reflex   will   be  excited  or   in- 
hibite.1    as  in  the  ease  of  one  animal  that   was  train...!  to  r.-sp..nd  by 
the  s.-crction  of  saliva  to  a  tuninji  fork  vibratinj:  at  100  per  second.     It 
V  .-  found  that  no  secreti..n  was  prodiu'e.!  by  a  tuninR  forlt  vibratni!.' 
ni  i04  or  at  W.    ^Mu.di  work  has  nls..  b.-.^ii  .lone  with  the  skin  recei)t..rs. 
Thus    wlien  a  --'iven  spot  of  skin  is  stimulate.l  cv.tv  time  that  food  is 
presented,  this  b.'cmes  an  active  sp..t    for  th..  c.n.litioned  r.^rtex.     At 
the  same  time  another  sp..t  may  be  stimulat.-l  so  .-.s  to  be  ass...-iated  liy 

the  animal  with  the  nonpresentati.m  of  f 1 :  it  is  a  conditioned  rofley. 

for  no  food,  and  is  a.ssociated  witli  the  absen.-e  of  salivary  secretion. 

Bv  eomparin-.'  the  resp.mscs  from  a.'tive  and  inactive  spots  wlien  both 
.nre'sti.nulat.Ml    either  .simultaneously   ..r   at    .-b'se    interv.nls.   much    can 
be  learned  cneernins  the  delicacy  of  appr.ciation  f..r  externa!  stimuli 
and  the  influence  of  the  inhibitory  ..n  the  ex.-itatory  process.     I'.ayliss 
cites  tl<e   followin<;  .-xperiment.     ALai-   a   series   of  si.ots  on   the  skin 
of  the  IcR  five  devices  are  arran-.'d   for   i.rodueinir   e.|ual   mechanical 
stimulati.ms  ..f  the  skin.     The  f.nir  up|.ermost  ..f  tliese  are  made  active 
spots  for  the  salivarv  r.Mb'N.  an.l  tli..  bnvest  ......  inaetive-that  is,  when- 
ever it   is  stimulat.'d  1...   fo...l   is  pr.sent..d.     L.-t   us  suppose  that   up..n 
a.lministcrintr  mclianical  stimuli  ..f  .-.lual  iiit..nsity  to  each  of  the  active 
f..ur  spots,  a  ce.tain  amount  of  saliva  is  i.r..duc...!  in  a  certain  time;  it 
now  the  inactive  spot   is  stimnlat.-l   an.l  th....  thirty  se.-.m.ls  later  .uie 
of  the  npp..rm..st  sj.ots.  ih..-.'  will  be  no  s..cr..tion.    The  previous  stiniu- 
lati.ui  of  the  inactive  spot  must  ..vi.b'ntly  hav..  cans..,!  an  inhibition  t..  be 
set  up  in  the  nerve  cent..rs  concrn...!  in  th."  v>-ih's.     This  inhib.ti..ii  ..nly 
.'ra.luallv  pass.'s  auav.  ,lisa,.p..a.ii.L'  fi.st   in  the  spot   fai'th.-st   rem-.v.-l 
from  that  made  ina.'tive.  but  it  may  take  s..v..ral  n.inut.'s  bcf...-e  all  th.' 
a.-tive  •spots  have  f,.a..(iuir..d  their  orijiinal  s.iisilivity. 

Th..  i.e.sis1.,.ee  .if  th,.  iidiibili,,n  PI...1. 1  by  slimulatinpr  the  inac- 
tive sp,.t  in  the  above  exi.,.rim..n«  in,li,.at,.s  an  in.|.,.i'tant  fact,.r  in  .m.u- 
necti.m  with  ',li..  pi'odiietion  .-f  .•,.n.liti,.nc.l  lellexe-;.  l',.r  .'xample.  an 
animal  can  be  train...!  t-  kn..w  that  in  a  c.tain  number  ,.f  minut..s  after 

the   sound    of   a    uiv..n   b..ll    f 1    will    b-.    iuvscnt..d    1,.   him:    the   .-omli- 

Ii,med  n.fl.'X  will  bec.me  ..stablish.'d  s,.  that  !..■  salivat.-s  at  .xactly 
the  san.e  tim.'  after  the  b.-ll  is  soui.d..d.  Som-'thinsr  must  be  sroins;  ou 
in   the   .•.■iit,.rs   durin-   this   time     som.thin-.'   inhibiti....'   the    reflex.      Tf 


I.    I 


(■(iNi'lTHiM  II    \SI 


1   NldSlirnoM  I)    F{l  Kl.l  XI  S 


85!) 


(hiring  tliis  iiitprval  of  iiiliitiilioii  some  I'tlicr  s.^iisory  >tiMiiiliis  is  app 


lied, 

il   uill  lie  iiixc'l\    tci  flit  sliiiit   llic  iiiliiliili.i'i;  in  otli.T  \\(ii<ls,  it  i)r.i«iii(M's 
ail  inhiliitioii  oT  inliiiiitinii.  so  tliat  tlir  .secretion  oi  sriiva  oceiirs. 

ilustiated 

llRllt 


Another  most  cMiious  eoinliination  oi  com 


lition 


stnnnli  is  i 


in   tlio  following'  experiment.     Snppi 


\i 


I'Xiimplc,   tiiat  a  {?ivei 


iiiil  soiin< 


1  an 


rli  sepaijiteiy  made  a  sliniiiliis  for  a 


intlitioiied  icIlcN. 


Ill  liiat  w  Inn  tliey  occur 


toLM'tlier  tlu'iT  is  no  reflex.     Supiiosc  now  lliat 


kiiile  one  of  these  active  s 


IS  a 


Iso  |ll 


eseii 


ted;  tlio   |-e.-<U 


tiimili   is  heinvr  presented,  tlie  other  stimiihm 
It  will  he  that  tlie  se<-ictioii  produced  by  tlic 


one  stimulus  will  stop.    Evidently,  a 


acting  together  they  <-anse 


inhiliition 


Uy  st\idying  the  coiiditioin 


d  reth 


Ithougli  each  is  in  itself  a  stimulus, 
it'ter  a  certain  part  of  the  cere- 


hral  cortex  has  liccn  rcniovt 


lisliiii 


■rtaiii 


1,  it  has  l)cen  t'otind  that  the  power  of  estali- 
s   Iiecouies    aliolishid,    while 


ids   of  conditi'-ned    re! 


that    t'nr  others   is  retained. 


1*1 


i  1* 


I   ':: 


I'l  • 


•  I   1 


•  1 

1 

i 

.h! 

«', 

! 

i ' 

|i 

!'■ 

(  IIAITKU   XCVIII 

Till-:   IIKIIIKK   I'lNt  TIONS  (•!•'  TIIK  (KKKI'.Wl  M    IN   -MAN; 

AIMIASIA 

■nu"  sliicly  ..I'  til.-  liiirli-'i'  fiiiii'ti.ms  ..f  tli.'  cvrcl.rum  l.'ii.ls  iis  to  111.-  ln.nlcr- 
laiiil  lM>i\v.'.'ri  |.li\>inlc,L',v  iiTi.l  |iMvli.il..i;y,  Imt  into  tliis  vii-t  an. I  ivhitiv.'ly 
uii.'Xi.loi-.'.l  li.1.1  w  Mil  ii.it  v.'iitiHv  h.'iv.  iiiili'-s  just  V.ir  .•n..ii'_'li  to  vMin  a 
Miilalil.'   xaiilair.-   I'.'iiit    rr.Mii    wlii.-li   to   uii.l.'i-Nlaii.l   tin'   iiatliol..i:y   of  the 

,Miii(liti.Mi  kiioun  iiN  .»/>/"/>/'/.*     As  wv  liav.'siMMi  fr ir  stii.li.'s  on  .•.•ivliral 

l,M-ali/ati.iii.  til,'  .■.Mvliriiiii  iiiiisl  !..•  iv-anl.'.!  as  a  -ival  s..nsoi-ini..t..r  van- 
L'lion.  «li.isi.  |-iiii.'ti..ii:il  a.'tivili.'s  ar.'  in.ii.-at.'.l  l.y  var'n.us  niovciiicnls. 
Til.'-.'  ni..v.'ni.'nts  may.  in  •.'.•n-'i-al.  i..-  rlassili..!  as  ol.J.M-tivc  in.li.-ations 
.■itiin-  of  f.vliML'  an.l  .■iiioli.m  ..r  .0  int.'lli'.;<'nc.-.  Altli.Mi'jii  liolli  .-iasscs  an' 
.■vi.l.'iil  in  all  animals,  it  is  paitiiMilarly  in  tli.'  .'as.'  ol'  man  tliat  tli.-  .'Vi- 
.l.n.'.'s  of  inl.'Uiv'.'nt  a.-1ivily  ar.'  .'sp.'.Mally  |.romin.'iil.  since  tli.'y  niclii.l.' 
..nMi.'iilati.in  an.l  tli.'  muscular  a.-tiviti.'s  iv,|uir.'.l  in  spoken  an.l  written 

laii'.'ua'.'.'.     Tl 1. 1\. 'Ml. Ills  tliat   .'spn'ss  emi,li..nal  .•.)n<litions  are  evolv.'.l 

i.arli.T  an.l  rr..m  low.r  plaie's  than  tlios.-  of  iiitell.'.-tilal  a.'tivity.  Tims, 
very  y..n:-  infants  ••niak.'  fa.'.'s"  wImmi  tli.'iv  is  ivas.m  1..  lu'li.'ve  tli.'y 
fe.'l  pain.  an.l.  as  lli.y  .lev.'l.ip.  tli.'ir  pow.T  .,f  .'xi-ressin-^'  .•in..ti.iii  is 
.'volv.'.l  l.'iiL'  l»'f.iiv  tli.'y  pr.-s.'iit  .'vi.l.'ii.'.'  .'f  int.'lliir.iit  motor  aelivity. 
an.l  still  l.iii'.'.r  li.'fon-  tli.y  .'an  arti.-iilate  wor.ls. 

Th.'  pli«'n..iii.'n..ii  of  liuman  psy.-lii.-  a.-tivity  wlii.-li  is  of  jrr.>at.'st  im- 
p.)rtaii.-.-  is  that  of  /.(/../."(-/'.  an.l  to  iiii'l.'rstan.l  lli,'  nature  .if  th.-  .•.■r.'hral 
inteL'rali.m  iv.|uir..l  to  iir.Hlu.'.'  it.  w.'  must   hrii-tly  .•.msi.ler  tli.-  .'.'rehral 

,„ >s,s    involv.a    in    th.'    iiit.'ll.'.Miial    .|.'v.'|..pm.'iit    ..f    tli.'    infant.      Th.' 

tirst  st.'p  ill  this  .l.'v.'l.ppm.'iit  is  th.'  st..rin'/  away  in  pn..ieetion  e.'nt.'rs  ..f 
m.'iiiori.'s  .if  th.'  s.'iisations  wlii.-li  th.'se  .•ent.Ts  have  ive.'iv.'.l.  F.ir  .'x- 
ampl.'.  wli.'ii  th.'  ehil.l  l.ioks.at  a  h.'ll.  ih-'iv  is  st..n',l  in  the  visual  .'.'iitiT  a 
meiii.iry  of  the  shape  of  th.  Iu'll.  an.l  wli.'i.  tli.>  h.'ll  m.iv.'s  s.,  as  to  pro.hi.'.' 
MPiin.l.  this  als.)  is  St.. re.;  a-  a  soiin.l  impr.-ssion  in  the  audit. .ry  .'ent.'i-. 
I.iki'uis.'.  when  h.'  t..u.'h.s  tie'  li.'ll  iiii|ir.'s>ioiis  of  its  har.lm'ss  an.l  snm.ith- 
ness  an.l  t.'iiip.-ratur.'  ar.'  si,.r.'.l  in  the  .'eiif.'rs  f.ir  .•ntam-ous  sensati.ms. 
At  lii'st  each  of  th.'sc  t!i<'!ii.ivy  i!iinr<'ssi..!!s  (..eeu)>i.'s  an  isolated  jiositioii ; 
but   later,  assoeiathm  tracts  .ip.'ii   u|>   hetw.H^ii   th.-ni,   s.i   that   the  calliii'^' 


•i'lir     'l-i-     ..f      I:..ll"ll'-     illlu',      i-     till. I.-     11.     Ilii>     .h.l|.l. 

cSiiO 


lIKilll.R   Fl  N(  TIONS   OK   TIIK    (I.HIMHrM    1\    MAS;    AI'MASIA 


■Mil 


forth  1)1'  one  iiii'iiiorv  imprrssinn  is  iissociatcil  witli  otlicrs,  iiiiil  tlif  child 
I'onii's  to  ln'  al)l('  to  associate  tlic  appfaraiicc  or  iiiiatrc  of  the  hell  with  a 
certain  soiiikI  ami  «  ith  cci'taiii  sensations  of  iiardiii'ss,  lotiiiulit y,  cti'.  This 
preliminary  use  of  oliscrvation  is  known  as  in  nriition.  it  involvfs  tlio 
fnsjfin  of  iliri'i't  sensations  as  well  as  their  correlation  witli  memory  im- 
pressions of  former  sensations.  The  numlicr  and  variety  of  the  latter 
called  into  activity  liy  a  particular  sensation  will  ulivionsly  vary  at  dif- 
ferent times.  On  seeinir  a  hell,  for  example,  a  I'hild  may  assoeiat<>  it  with 
sonnd  on  one  occasion,  and  on  the  next  with  tlie  fieliriir  of  the  hell.  On 
account  of  this  diU'erencc  in  the  detail  of  the  method  of  association,  it  is 
evident  that  pei'ception  iinist  he  a  prodili  t  of  cerehral  integration  I'atlier 
than  one  dcpeniliie.'  on  iiic)noi-y  impri'ssions  stored  in  the  isolate  I  eentei's. 
It  is  a  complicateil  pr'occss  with  an  infinite  vai'ii'ty  of  possiliilitcs  as  to  the 
exact  wa.\'  in  which   it   is  integrated  on  each  occasion. 

The  act  of  jicrccptioM.  Iiowi'ver.  liccomcs  considcrahly  simplilicd  in  the 
hiirher  animals  hy  the  layiiiL'  down  of  slmrl-riil  iinll's  'if  ns^oiiiifioii. 
These  are  formed  lii'st  id"  all  .vith  the  auditory  center,  in  which  the  memor.\- 
impression  of  an  articulated  sound  represent iuL'  the  oliject  for  example, 
the  word  '•licH"  is  stored  away.  The  child  comes  to  learn  that  this  par- 
ticular word  is  to  he  assoc'iated  with  the  memory  impressions  it  has  stored 
awa.v  of  the  sound,  the  siuht.  and  the  feeliic  of  the  1"11.  Similar  shortcut 
paths  later  lieconu'  developed  in  eoimectioii  with  the  visual  eriiters.  where 
a  c-ertain  syiiihol.  like  the  word  ■■hell.''  is  ju'escMiteil  to  the  child  as  si'j-ni- 
fviiiL'  all  the  other  attrihutes  of  hell.  In  its  most  hi'jlily  developed  form, 
tl, el.  fore,  perception  uia\  he  deserihed  as  the  ;ict  o|'  c.dliii'.r  up  one  nr 
more  seiisorimemorial   imayi  s  wlieii  a  name  is  seen  or  heard. 

Ilaviiii.'  acipiired  the  ahilit.v  to  intci^rate  sensorimemorial  iniprcssiims 
in  the  ahove  descrihi'd  manner,  the  c'hild  next  learns  to  intetrrate  the  motor 
centers  eoneerned  in  the  control  of  the  articiiiatoiy  apparatus  sn  as  to 
produce  a  sound.  This  somiil  is  the  word  indieatiliir  the  nhicct  involved 
in  the  iiitctrratinir  process.  It  is  the  inteL'rati<in  necessiiry  to  produce  the 
soinid   wliii'h  symholi/es  the  jtarticiilar  oh.je<-t. 

AVhen    the   power  of  miderstandinir  and    iiroclucinir   lani.'uai.'e   has   hecn 
accpiircd.    the   crowninir    jiroeess   of    intellectual    devclopineiil      thi'    forma 
tion  of  a  roiifiiit.  or  crencral   notion     hecimies  evolved.     Thus,  the  evolu- 
tion of  a  treneral  name  will  include  a  inimher  of  partii-iilar  oh.iects  or  acts. 
■'This  process  of  e(Uiception    invohcs  the    rc\  ivitication    of   iniiiierous  sei:- 

^oiinii'iiiorial   imairi's  which   present   con m   points  of  siniilarit.\  "'      i  Mol- 

l(Ui\  Tt  is  relatively  ii  -imp'--  p.i->ccss  f.o-  su.-U  L-ctiera!  (ihjeds  as  a'.iimal, 
man,  huildin'^,  hut  hecomes  \ci-y  complex  for  sucli  ahstraet  coiKM'pts  as 
heaviness,  heanty,  etc.     It   is  ohviously  a  process  to  which  no  one  cerehral 


]     * 


<Ml 


Iff 

~ 

'v. 

1 

i 

•    1 

li  ?li 


I       !  I 


862 


TIIH    rKNTllAI.    M  KVOI  S   S\STi:M 


,,,„,,,  ,a„  1,.-  Mssiu'M.Ml.     Tli.  .mtu7.nl   Muu.if..station  -f  llio  .•.,M.T,.ti..i.   is 
spok.ii  or  writ t I'll  laii'/iiiij:«'. 

La, ,..'(•  .'onsiMv  tli.-ivl-nn..  i xti-.m-ly  .Mnni.l.v  ^vimIm.!...  syst-n., 

involvinir    Viiri-ms    ,...„t,.rs    a.,.1    Mssn..i,.tin„    tra.^ts    in    tlio    r.-r.O.r.nn     a,..l 

,a,,al,l..  of  ,.n  aln.M   i.iti.ut,.  .l.ur f  .l.v.-l..,.nu-..t   l.v  Iho  laying'  down 

„,■„,,„.  .v,nl.oli.  sN>1.n...     Lanu'ua..',-.   i.-l 1.  l'0.-"".-s  tl.o  insln..,.-..t    nl 

,|,ou«l.t  pra.ti.Mllv  all  of  tlir  Imu'Iht  iul.li.vtual  ,.nM-'s...H  !>.•.,.>.'  .lopon.ln.. 
„„  its  ovulMti.m.  I.,  this  ..onn.rii,-,,  it  is  iMtrr-stin-  t..  n.M.  tliat  a  S-'roal 
,„„„,,..,  „r  i,„livi.luaN.  ..s,„vially  tlH.>..  wlio  ,1,.  M.,t  n.a.l.  .lq.n.,1  .m  IIm' 
s,.„M.  ,.!•  iH.ari,,..'  fnr  tl,..  a-.,uisitic.M  ut  ll„.  i,n,.n'ssions  ro.,u,f.-.l  for  tho-r 
psy..|,i..  .l.'v..|o,„ t.   win!,-  utlins  .l.-p-M.l  on   th-  mm.so  ..f  s,j:l,t    lor  tl,. 

""\i  ll'lTrimv  -lilV.Mvnt   tMH-s  of  ,-..nt.T  an-  invnlv.l   in  tl,o  int.>irn.li..n 

„r  I,, „ani-lv.  au.litory.  vi-nal.  .•l,iioirfapl.i<-.  an.l  aitn-ulatory.    \\.' 

,„av  .all  thp-..  -'wcnl  n.nt.Ts,--  an.l  w.'  must  assni.io  that  tl.-y  1...  n.-ar  to 

,|„:  a.i.litorv.   visnal   an.l   t-'-n-ral   s-ns..ry   pn..i.-ti...l   ans.s  .,|    tl .ft.'x. 

■r.,  „,„l..,stan.l  an.l  to  U-  Mv  t,.  |.n..ln..-  sp.-kon  an.l  Nvr.tt.-n  lanirua-.^.  H 

i,  , .sarv  tl.at   all  tl.-.  fonr  wonl  ■•.•nt.'fs  pafti.-ipat.^  thnni^'li  ass.,..,a^ 

,i„„  „,,-ts.'  altlMm.'l,  tiM.  anin..'  of  a  v..nl  n.ay  1,-  p.M iv.'.l  w.tlmnt  all 

of  til. 'Ml  iM'in'-'  invoK.'il. 


*    ill:' 


H\     ! 


.        i        I 


PSYCHOPATHOLOOICAL  APPLICATIONS 

1„  ,l„.  s1u.lv  of  ni.ntal  .lis..iis..s  tli.'  in..st  important  .•on.-lusi.ui  ul.i.'li 
„..  ,.;,u  .Iraw  Vn.n,  tl,.-  al...v.-  faw-  is  that  lan^'uai:.-  is  .•s.s,.ntn.lly  a  sym- 
,,„l|,.  „„-.-l.anism  f..r  tla-  int.L'niti..n  ..f  s.-nvriun-morial  nnafres.  It  is 
tl,on-f..n-  til.-  svml...r,..  sysl.m  ..f  tin-  int.-ral.-.l  pro.-.-ss.s  ..f  tlu-  l-ram  ;  it 
i.  ,|„.  M-rvant  ,.f  thou-ht.  \VI,.-n,  as  is  oft.^n  tl,.-  .-as.-,  lan-ua-.-  ,s  us.-.l 
without  tl,..  pn.p.T  ..V.  .vis,,  of  tl,o,...'l,t.  ,t  l...,-ou,.-s  n,.-n.|y  an  aut..mat,,. 
,ir-,ir       \  i.ra.-ti..al  .l..l,u-tioi,   tVo,,,  th.s.-  fa.-ls  is  that   any  ..onsnl.-ral.l.. 

,1,,-M lu.-nt    »(  th-   lan..'uauv   ......hanism    must    i ssarily    inv..lv.-   s.,.,..- 

i„„.,-f..n.n...^  with  tl„.  .•onipli..al...l  pi s.s.-s  ..f  ass...-iati..n  that  v"  t..  mak.' 

\ip   tl,.'   psy.-hi.-    fnn.tioH. 

Th.-sc  .•.u,si.l.-rati..ns  naturally  l.-a.l  us  to  tho  subj.^ot  of  aphakia.     U  has 

1 „  usual   t,.  .listin.^uish   tin-....   vari..ti..s  of  this;   nanu.ly.   .,iot..r  aphas,,-,. 

M-nsorv  aphasia,  an.l  anarthria.  In  n,olor  aphasia  th.-  ,mti.-nt.  alth.-m^h 
1,..  un.h  rstan.ls  what  is  sai-l  to  hin,.  is  unahl..  to  sp.ak.  an.l  th..  i..toll.-.-tual 
p„w..,.s  an-  litth-.  if  at  all.  impain.!.  in  s.-nsory  aphasia  sp.wli  is  ,M.ss.hh. 
i„  a  „„.,...  or  l.-ss  int..|liuil.l..  .„ain,..r.  hut  tli-n-  is  a  .listin.-t  impain„..i,t  ..t 
i„t..llh'.-n,..^     In  anarthrin.  or  snh.M,rti.-al  aphasia,  th.^  only  .lisah.hly  is  Iht- 

loss  orimpairm-nt  .,f  tho  p..w.M-  of  arti.-iilat..  sp. li  h.-.-aus,-  .,f  son,.-  h-s...n 

..Nistintr   ill   tho  .•.-nt.r  .•n..r.linatin-.'   tho  l-.wor   n.-iu-ons  .•on.-.-rnt-.l    in   tho 


IIKIIIKK    n  S<  TIONS   or   TIIK   riHIJIKIM    IN    MAN;    AI'IIASI.V 


8fi:» 


nixvciiiiiits  ol"  tliu  larjntjt'lil  hiuI  Umguc  muscles.  Piorrc  Mnrit«,  uh  u 
i-.'Milt  (if  \iv\  i'\\vns\\v  i'\|.frii'ii'-c  ill  Paris,  lias  shown  that  this  claswifl. 
cation  is  iiii.iii-.titic(l.  Ill-  iiuiiiilaiiis  that  thorp  is  only  one  true  form  of 
apha.sia.  ami  that  Mich  a  thiiii:  as  iniiv  motor  aphiutia  as  ahovc  (hfiiicil 
floes  not  exist,  the  eotulilioii  iieiii;;  iiivariaMv  aeeoni|mnie(l  liy  inlelleetual 
impairment. 

Marie  points  out  that  the  various  elaims  that  aphasia  may  exist  without 
intolleetual  impairment  have  tieen  nunh-  without  siitlii-ient  investiiration  of 
the  iiitelli'elii.il  status  of  the  jiatii'iit.  He  shows  tliat  many  patients  siif- 
forintr  from  aphasia  if  askeil  to  ilo  onliiiarv  thiiiirs,  siieh  as  coti^'h  or  spit 
or  raise  the  hainl.  can  <lo  Iheiii  as  well  as  a  normal  imliviclual,  hut  that 
tliese  after  all  are  very  eriide  aets  in  the  onliiiary  performanees  of  a  normal 
individual.  Tr)  test  the  intellectual  powers  it  is  necessary  to  reipiire  the 
patient  to  |m  rform  ads  which  entail  a  considcrahle  amount  of  eerphral 
inteL'ration.  We  must  ask  him  to  |)crform  smne  seipienee  of  events  siieli 
as  walkinsr  several  times  in  one  direction,  then  in  another,  touchiiii:  cer- 
tain olijei'ts.  etc..  or  lieltcr  still  we  should  ohserve  the  patient  closely  in  his 
husiness  transactions  and  everyday  routine  of  life  to  sec  whether  he  docs 
thinsrs  exactly  as  he  did  tli.'iii  hct'ore  It  is  always  possihlc  hy  such  tests 
to  show  that  in  aphasia  tin'  mcnliil  powers  have  licconie  distinctly  dc 
prcciatcd. 

The  portion  of  the  I'crehr.d  cortex  atVccted   in   aphasia   is  always  in  the 

iicitrlihorl 1  of  the  uncalled  area  of  Wernicke,  which  is  closely  related  to 

the  visual  and  aiiilitory  centers.  In  makiiiLr  this  swecpim:  conclusion. 
Marie  admits  that  cases  of  pure  word  hlindiicss  hut  not  of  word-deafness 
may  exist  ;  that  is.  a  patient  still  retainiuL'  his  intellectual  powers  may  lose 
his  ahility  to  interpret  correctly  what  he  sees,  althouirh  In-  isin  still  interpret 
aceiiratel\    what  he  hears. 

This  conclusion  conforms  exactly  with  tlio.e  of  the  psychophysioloirists 
reirardini.'  the  dilTcrein'c  in  the  laiiL'uairo  mechanisms  of  educated  and  un- 
educated persons.  l,ani.'iiaL'e  is  learned  throuj.'h  the  sense  of  hearinsr.  and 
it  is  only  hy  later  eilucation  that  more  is  learned  hy  the  sense  of  sifflit ; 
that  is  to  say.  a  pei-MMi  learns  to  read  only  after  he  has  learned  to  under- 
stand spoken  laiiL'iiai-'e,  Wordhlindness  may  therefore  occur  as  a  pure 
s\  inptoni.  ami  is  less  likely  than  \v<ud  deafness  to  he  associated  with  nli- 
nornial  inteiri-ntivc  functions  of  the  cerchrum.  Word-deafness  liowever  de- 
p.Mids  niton  a  lesion  involving'  the  auditory  center:  it  necessarily  moans 
ilisturhancc  in  the  asso.-iation  functions  of  the  eerehnim,  and  is  always 
accompained  hy  a  certain  amount  of  mental  deraiiffoment. 

Tti  coiTrrlH>ratioii  of  these  facts  may  he  cited  the  well-known  fact  tluit 
a  (h^af mute  is  mentally  far  inferior  to  one  tliat  is  eonpfpnitally  Mind, 
r.oss  of  hearinsr  loads  to  more  serious  ceroliral  disahility  than  loss  of  siprht. 


il  'H I  ' 


s\- 


'     It 

r 


!-('ilri 


\-  ' 

i. 

If 

1 

t| 

i 

■i  *■ 

^ 

1 
1 

I  ! 


*        I 


|i| 


Ml 


Tlir.   rlNTHAI.    M  RVdlS   SVMTKM 


Ti)  <iiiotc  Holtoii  iitfiiin,  •III  Mii'h  ••iiMs  .l.-afiifss  i-*  tli.Tcforo  a  morn  nt-rum^ 
tlpprivatii.li  than  IiIIikIik'ns.  as.  I'..r  tin-  .•volution  of  tlif  Iniu-lional  activity 
of  the  corcliriMii.  an  fiitir''lv  lu'vv  .|.'Vfl<>|iiin'iit  »(  a>»sofiatiimal  »|>licri'H  tn 

r,.pl, thov  iioiinally  .niplov.,!   for  amlitorv  hihI  s|.ok.'ii   lanirnairf  lias 

to  li.'  a.'Muir.-.l.  In  tlir  .'as.'  of  .•ont'.'iiilal  or  .'arlv  .i,.,|uiiv.l  l.lin.lii(".x.  m, 
the  oIli.T  liaixl.  till'  .•onipl.A  >|.li.  iv  of  laiiiriiaL'c.  with  all   its  psvrhic  .-(.iii 

,, „|H.  ran   1 iii|ilo>.<l   III  a  iMTffflly  normal   nianii'-r  an<l  almost   .x 

a.'tly  as  it  is  luoiit'lit  into  us."  in  tlif  .-a-.'  of  p.-rsoiiH  vvl itli.T  f.'ii.l  lan 

write' 

It  woiil.l  1m-  Ih.voii.I  111."  scop.'  of  this  work  to  jro  int..  tli.>  .•lini.-al  aii.i 
pnth..l..t'i.'al  .'vi-hii.--  iip.>n  «lii.'li  .Mali.-  Las.-s  his  far-r.-a.-hin-.'  .■on.-liisi..iis 
Siitli.M-  it  to  sii\  that  it  is  .htinit.'ly  shown  thai  ih.'  '.I'l  .•ont.'iil  i..ii  ..f  ltn..-a 

tiiHl   a  s| ial  sp.'.-i'h  .•.-iit.-r  .-visls,   is  .mIuvIn    mijilslili.-.l   hy   Ih.-   fa.-ls  ol 

i-lini.-al  aii.l   patholoui.-al   .-MM-ri-'ii.-.-.      I'.ro.-a.   it    will   h.'  r.-iii.liil»-f.',l.  ■■on 

tc-n.h-il  that  inol.ir  aphasia  is  :il«a>s  .lii.-  1..  .l.-s1ni.-1  iv.-  pi ss.->  .M-.-iirrih-.' 

ill  th.-  lower  portion  of  th.-  as.i-n.lini.'  frontal  .■.involiiti m  th.-  I. -ft  snh-, 

and  h.-  .-on.'ln.h.l  that  this  porti.in  «(  lh.>  .•.-r.-hriini  npr.vnls  tin-  sp.v.-h 
.•.-nlir.     Marie  has  shown,  h.iw.vrr.  that  a  pali.-nt   iiia\   sh..w  .listin.-t   .-v  i 

,|,.,„ f  aphasia   without   any   hsii.ii    involviii!.'   this  s.i  .-alh-.l    I'-i a   ar.-a. 

an.l.  on  th.-  ..Ih.-r  haii.l.  that  .-as.-s  ii.it   infr..|ii.nlly  ur  in  whi.-h  this  is 

,.,.niph-li-iy  ihstroxr.i  wilh-iul  any  .-vi.l >(  aphasia.     I  iiip.ntant  tlion-.-h 

this  .lis,-..v.-ry  -if  th.-  ina.Miira.y  of  r.r.M^'s  .-.in-lnsi.ai  is.  I..\  far  th.-  in.'si 
important  .-..n.-liision  whi.h  w.-  may  .Ir.nv  from  Mari.-'s  work  is  that.  :  in.-' 
laiM_'iia-.'.'  is  a  pn.,in.-l  of  an  .-\ti-n.l.'.l  inl.-i:rali.in  .-f  ii,ipivssi..ns  and 
m.-m..ri.-s  ston-.l  in  .lil1Vr.-nt  parts  of  tin-  .-.-r.-hruiu.  it  w  not  s.,  lik.-ly  to  1m- 
int.-rlVr.-il  with  hy  .1.-st  rii.-l  i.ni  of   uiv  oin  »f  th.>  .-.-nt.-rs  as  if  is  l.y  .h>strn.- 

tion  of  th.-  paths  which  .-oni t  th.-  c-nt.-rs  with  -ai.-  aiioth.-r.     As  a  matt.-r 

of  fa.-t,  Mari.-  has  shown  that  in  .-as.s  nf  a|>hasia  tin-  l.-sion  is  ii.-arly 
always  lo.-at.-.l   in   th.-  .■..iirs.-  ..f   th.-   pathway   .•..nn.-.-tini:   the   visual   and 

auditory  ,-.-nti-rs  with  tl th.r  .-.lit.  is  ..f  th.-  .-.i  vl.nim  ;  it  li.-s  aronn.l  tin- 

iipp.-r  .-11.1  ..f  th.'  li^siMv  of  Sylvius  in  llu-  r.-i:i.in  wlii.-h  in  pr.-vi..iis  y.-ars 
had  h.'-'ii  .-onsi.l.-iv.l  parti.-iilarly  ass...-iat.-.l  with  th.-  .•oii.liti.m  kin.wn  as 
s.-nsory  ai.hasia.  Th..s.-  int.-ri-stf-.l  in  this  snl.j.M-t  sli.ad.l  .-oiisnlt  Hollon's 
article. 


(  IIAITKU   .\«  1\ 


KINfTluNS  (IK   IIIK  (  KHKIIKI.M  M 


III  (iiir  (lisiMissioli  of  rrtlcx  iirlioii   we  liavc  *ii   fur  roiisiilcri'd  "Illy   tlmsc 

!•( ptiiis  rniiiiiic  I'i'i'lii  fill'  i'\ti'ri<ir  1(1'  the  Imdy,  iilllinii'.'h   \\r  lin\c  ' kv 

iii/i'il  lliiit  11  I'liiisidiTiilili'  iiiiiiiIkt  lit'  tlio  alTiTi'iif  iitTvc  rmits  rcmtiiiii  tiliii's 
(•"iiiini.'  I'iniii  rci'i'iitors  siliiiiti'il  in  tlic  iiiiim-Ios.  tlio  ti'iiilniis  iiinl  tin'  jiiiiits. 
iiiiil  ciilli'd  |ir(i|iriciiM'pt(irs  lnTiiiiNO  tlicy  ri's|i(iii(l  imt  to  cluiiiL'fs  in  llw 
riivifciiiiiiciil  Imt  III  iiltcralioiis  in  tin'  lioily  itsi'If.  AVo  Imvi'  situ  that  the 
|irii|iriiH'i'|itiir><  rmisisf  Ntniftiiriilly  nf  nnisclr  spinilli's  anil  of  lln'  ihtvc 
(Milliners  in  tin'  ti'iiiliiiis  iiihI  jitratui'nts  ami  syiidvitil  iiii'iiilirani-i.  'I'ln-y  an- 
ri'c'ppfiii's  that  arr  atfiinril  to  ri'spimil  tn  ilitrorrinM's  in  ti'iisimi  raii-i'il  oitlior 
li>  liuij.'iii;,'  nl'  till'  iiiiisrifs  Id-  liy  si I'l'tcliint:  "f  tlir  liliiTs  id'  Icinlciiis  and 
liiriiiiii'iits. 

Till'  iiiipniM's  an-  transinitti'd  in  Ihi'  >piiial  ronl.  i'iiImt  hy  tlir  puslcrioi' 
foliiihiis  111-  \>y  till'  lati'i-al  riTrlirllar  trarls.  'I'Iium'  travi'liii!.'  Iiy  thi-  pos- 
ti'i'icii'  I'liiiiiiins  an'  si'nt  mainly  In  tin'  ri-ri'liral  rniti'x  nl"  the  uppositr  sidi. 
whi'ii'as  llidM'  in  tlir  rrl'i'lii'llar  trai-ts  cnttT  tin'  ci'i'i'lii'lliiiii  hy  tin'  iiit'rrini' 
pru'liich's  111' till' saiiii'  sidi'  TIii'  I'i'i'cliial  iiiipiilM's  I'liinn'ct  with  ni'iiniiis. 
whii'h  ti'aii^iiiit  tin'  iiiipiiisc  hark  a'/aiii  to  tin-  n'ri'lu'llimi  nt"  thr  nppositi' 
siih'.  so  that  nltitiiati'ly  thr  ciTi'lu'llar  nirti'X  is  I'linni'i'ti'd  with  thi'  spinal 
I'lii'd  III'  tin'  saini'  sidi'  i-itlnT  dircrlly  nr  iiidirrrtly  thrcniirh  tin"  ccirtiral 
c'cirt.'X.  Thi'sp  anatiiiiiii'  facts  indirati'  in  a  ir"'"'"'';'!  "ny  'li'''  ^^'''  ""i\ 
I'xpi'i't  till'  fnnction  "f  tln>  ciTi'lii'lliiin  to  ho  that  of  the  rliiif  in'rvc  I'ciitiT 
c'liiii'i'nn'd  in  tho  iiiti'jrration  of  tin'  in'oprioi'rptivo  impulses  oriL'inatid  hy 
till'  I'lPMilifioii  of  contrai'tion  or  rrlaxation  of  the  dilffrcnt  irroups  of 
miisclos  in  till'  hiidy,  and  liy  the  amoiiiit  of  ii'iision  oxistinsr  in  tin'  various 
tendons,  liirameiifs  and  other  iiieniliranes  surroundiiii:  the  joints. 

Kxperimental  iiivestiiration  has  .justified  those  expeetatioiis.  Tho  I'e- 
moval  (if  the  entire  eereliellum  an  operation  whieli  has  usually  lioen 
performed  on  hirds.  partieularly  piproons.  lieeause  of  tho  ease  with  which 
it  eaii  lie  done  in  tliOM'  animals— leads  immediately  to  a  rondition  in  wliieh 
museular  aetivity  is  entirely  uncontrolled.  A  pit'con  after  this  oporation 
flies  alioiit  ill  an  ini'oordinato  way.  tiirninir  summersaults.  ilasliiii<r  itself 
airainst  the  walls  of  its  I'hiiinlier  and  nltiiiialely  after  constant  futile  move 
inents,  exhaiistiiijr  itself.  If  one  eereliellar  lolie  is  removed,  the  hody  when 
at  rest  is  curved  toward  the  side  of  the  lesion      -id  the  niovomonts  of  the 

Sfi.'> 


If 


n 


Lii 


8«;(! 


Tin;  (i;ntkai.  nkuvois  systi:m 


M„i,nnl  ouusc  it  to  fall  in  tliat  direction.  A  similar  .■x|HTiiiu'nt  with 
,l,..'s  Yields  like  results,  l.ut  the  oi>erati..ii  is  "f  .-(.iirse  .■..iisi.lcrahly  more 
dirtieillt  In  mail,  a  d.Mructive  tumor  of  the  .•erel.ellnm  prodm-es  a  eoti- 
ditio,,  known  as  •■.vrehellar  ataxy,"  in  whieh  the  ,.atient  moves  jus 
limbs  in  a  very  ineoor.linate  fashion;  ho  stantrers.  is  ui.eertain  in  Ins 
"ait  and  behaves  in  jniieral  very  like  a  drunken  man. 
'   .Mthou^'h    these    immediate    e(Te.-ts    of    eerohellar    extirpation    indicate 


I,.      ■    ■       |-„oi,„,„t,    ali.r    .U-ttuclion    of    the    ccrcLrllum    in    a    .log;      «.    l.t(..r.-    the    o|,er»tioii . 
^:  fmrr  da,  .!,';■.;"?.     ..V.  .lays  after;   ./.  a  „u,„.h  after;  r,  .«o  mon.h,  after.     (I-mm  I.uc.ni.) 

rlearly  thai  this  ori.Mii  hits  in  .h.  «itli  the  .M.iiln.l  «(  mnseular  uiovenients. 
y.t    tiif   restdts  ar.'   pnthahly   not   primarily  .jepen.lent   on   the  ahlatitm. 

"l,i,t   ,.;,ther  nil  il nnilitiuiis  of  irritati(m  which  are  set  up  as  a  result  of 

il,f  t.peration,  ami   whieh   pn.hahly  affeet   the  ecrehellar   pe.hineles.     At 
least,  sueh  is  the  view  that  Ijieiani,  one  of  the  greatest  iiivestitrators  in  this 

li.-M.  h.is  M.i.ipt.Ml  1 ausf  of  the  fact  that,  if  the  ••uiinial  is  k.'pt  ali\e  for 

>unieifnt  time  so  that  the  symptoms  of  irritath.n  disappear,  they  beeome 


ITNCTIONS   W   Tin;    CKKKHlll.I.l'M 


867 


n-pliu'i'd  liy  tlidsf  of  an  cnfiicly  (lilV.'iciit  iiaturc.  Tlic  pi-reon  may  reac- 
(juirc  tlio  piiwor  (if  Myintr  Mrai-rlit,  or  and  ttiis  is  particularly  important  - 
tlic  <lo<r  may  n'ai'i|uirc  tlif  ])o\v<t  of  appari'iitly  normal  pro-rrossion,  al- 
tlioni.'li,  if  its  nnisi'iilar  niovrnifnts  an-  carcfnlly  cxamim'd  tiy  physioloxif 
mi'tlioils.  it  will  Im'  foUM.l  that  at  l.'ast  tlinv  cliaiifrcs  liavo  dcvclopod  as  a 
lato  result  of  till-  extirpation:  namely,  a  weakness  of  eontraetion,  a  tremor 
during'  the  eoirtraetion,  and  a  want  of  tone  wlieii  at  rest.  These  condi- 
tions liave  lieen  <'alled  iisIIk  iiin,  nhiiiin  and  iislii.sin.  lespeetively.  On  su- 
perficial examination  it  may  often  he  dirtieult  to  make  out  these  three  con- 
ditions, lint  they  can  readily  he  oliMrved  in  animals  in  which  tli rehellar 

extirpation  lias  heen  performed  on  si<le.  so  that  the  al. normal  may  he 

compared  with  the  normal  side,  in  a  do^'  that  has  had  one  cerehellar 
hemisphere  removed  some  time  previously,  the  muscles  on  the  correspond- 
in".'  side  arc  so  mui-h  weaker  than  those  on  the  opposite  side  that  the 
animal,  in  order  to  retain  his  ccpiilihrinin.  has  to  prop  himself  up  either 

hv  leaning'  aj-'ainst  whatever  oh.ji'ct   nwiy   1 onvenieiit.  or  hy  extendin;.' 

his  le^'s  so  as  to  increase  his  hase  of  support.  In  other  words,  he  constantly 
tends  to  fall  to  the  side  of  the  lesion.  Imt  tries  to  prevent  this  either  hy 
increased  niuscnlar  efi">rt  or  hy  takin-r  a.ivanta<;e  of  artificial  supiwt. 
The  elTect  which  this  ■  keninpr  has  on  his  -rait  can  he  very  clearly  demon- 
strated hy  comparinj.'  the  footprints  produced  hy  the  normal  with  those  of 
the  ahnormal  side,  these  footprints  in<r  ohtained  hy  nuikin;.'  the  animal 
trot  alon-r  a  i>ieci'  of  ^rla/ed  paper  hlaekened  with  a  carhon  de|iosit  as  in 
takinir  trai-intrs  i  Fiir.  '122"'. 


Localization  of  Function  in  the  Cerebellum 

Althou-ih  these  facts  in  themselves  would  tend  to  indicate  a  certain  (h' 
jrrco  of  liiciillzdliim  of  fiiiitlliiii  in  the  eerehelhim.  or  at  least  that  certain 
parts  of  the  cerehellar  cortex  have  to  do  with  certain  }.'rou|)s  of  muscles, 
yet  for  many  years  it  was  consid.red  that  the  ccrehcllnm  did  not  show  in 
any  marked  dejrree  tiie  same  kind  of  localization  that  we  find  in  the  cere- 
hral  cortex.  One  cause  for  the  hackward  .state  of  our  knowledge  concern- 
ing cerehellar  localization  is  that,  unlike  the  cerehrnni.  its  cortex  is 
practically  inexcitahle.  In  recent  years,  however,  on  account  partly  of 
.inatciiii;.'  ,ind  partly  of  experiMieiit.il  and  cjinical  work  a  lii^rh  de._'i-ec  of 
localization  has  heen  I'oimd  to  exist  in  the  cerehellum.  From  the  auatomii- 
point  of  view  it  lias  heen  found  that  in  certain  v'"""!'^  "•"  animals,  such  as 
the   un^idata.   the   postcro midial   hihidc  of  the  ci'ieliclluin    is   vi'r\    laiirc 


whereas  the  lohuli  ansiformes  arc  small.     T 
the  opposite  ohtaiiis.  tlie  loh-i1i  ansiformes  hi 


n  another  u'roup.  the  carnivora. 
•..'leatly  developi'd  a"d  the 


lostero  mi 


dial   lohidc  (piite  small 


i  .11 


i  -1 


I     i 


ill 

ff 
iffl 


\P' 


I:':"'!' 


i  I 


it   I         Mi. 


■'I 


IIP! 


S68 


riii:  t-KNTKAi.  Nruvors  systkm 


Hy  stmlviii}.'  iIk-m'  developmental  differences  in  ivlaticnsliip  to  tlif  a<'iivi 
ti.'sof  th/iiuiMMiliir  syM.'in.  r.olk  Mi--.'Mr,l  iliiit  iiKivcin.'iit  of  those  rrgions 
of  llio  ImhIv  wliicli  iiiv  iitT.vtiMl  l.y  iiiiis.'l.^  LToiips  on  Loth  >i.l<'s'    for  rxainpl.'. 
tlio  lioiid.  n.vk  or  trunk     would  lie  ivpn-M-ntrd  on  tlic  n'lvlM.lliir  .•ortcx  l.> 

anunpair.'d. t.T-   tliat  is.; ntn .•upviii!:  a  mid.ll.'  position     and  tliat 

tills  would  1m.  .•apaM..  of  ..nitimsIml'  an  intlumc.-  .•(piallv  upon  tlu-  inusrl.v 
of  liotli  siili's.  Mov.iu.'uts  of  tiio  liiul>s  would  ivipiiiv  an  rntir.ly  ditT.'ivni 
tvp..  of  coordination,  -incr  tli.'y  ar.'  not  aci-ustoiii.'d  to  a.^t  t..-i"tlicr,  unlcs. 
fur  .-..rtain  niovuiculs.  as  walkin-     Has.-d  on  llicsc  lli.M.rctic  cousid.MMtion^ 

I'.olk  foiMid  a  d.'tiuit )rrcspondcMiM.  to  exist  Intwoon  tiie  variations  in  tli^ 

dov.'Iopnicnt  of  rrrtain  .MMvlM.llMr  lol.ul.^s  and  tlic  funrtional  iinporta of 

.•.•rtain  unis.^l.-  .jiou|.s.  and  tli.>  ■.'-•n-'ral  c.n.-lusions  d.  clu.'ii.l.'  from  liis  an. I 


l.,u  ;       iii..»..i,.m<    1"    r,ii.-'iit    r.-.|..vtr..K     .,    v.ilia:    m.  «     ■•(    liii'    Ml     li.>ll     ■•"■!    •>    'i"'-'' 

,,,  l;,,rK.  >V<:r<  l.1...t-K^,.|l,s  -I  -i.MiUius  tr.ini  11^.'  Aiuilun-.i.al  .Mn„-.im.  \\v-„Tn  W. -rv,-  M,-.|„.,l 
>,  li,...l,  I      .  1-r-m    |l.ivi.|«..n    I'.l.i.  I,  i 

(•orr.'lati'«l  work  tiia\  he  sniiiinrd  np  as  follows  i.'f.  Uavidson  l!la.-k"  '  :    Tlii' 

lolins  antcrioi-  .-.  r.'lMlli    s.c  [■'iy-.  'J-J:!    .•oniains  tli ntcrs  t".>r  the  .•o.>rdi 

nati.ui  of  till-  nnisi'li'  -groups  .if  tii.'  iica.l  i.ys.  toii<,'Uf,  nius.^lcs  of  mastica- 
tion. nius,-ks  of  .•\prcssi.in\  an.l  'if  tli.^  larynx  and  pharynx.  The  lohus  sim- 
plex I'ontains  .•.■liters  for  tli.' rdiiiation  of  tli.^  musel.'s  of  tho  neck.     Tlic 

Inhulns  iiie.lianns  posterior  .•ontains  the  iinpair.'.l  .-enti-rs  for  tho  syn.'l-frM- 
ni.iv.^m.-nts  r.^.piired  U\  iUr  riirht  ami  hft  .■xtr.'iiiiti.'s  for  the  purp.is.^s 
.if  iiroirression  <  Hi  th.'  oili.^r  haml  pair.'.l  .•.nli^rs  f.>r  th.'  .■xir.'imtii's 
th.is..  .-ent.'rs  that  hav.'  to  -lo  with  th.'  iii.l.-p.n.l.'Ut  inov.-im'nts  of  ca.-h  linili 
..f  the  saiii..  si.h-  .,f  th..  l..Mly  ar.'  I.i.-at.-.l  in  the  h.linli  ansiforiiios  ct  para- 
modiani   M-nis  |.rimnin  ami  .•riis  s.vundiini  > .     Tho  ..entiM-s  for  the  rost  ot 


m 


KiNcTioNs  or  Tin;  ci  ui:i'.i:i.i.rM 


Ht;i» 


flu;  trunk  iiiul  tail  rc^rioii  arc  locatcil  in  tlu'  rciiiaindcr  of  tlie  i-cri'lujllum. 
These  eonelusiiins  are  the  hasis  ut'  thi'  aeciiniiianyinj.'  map  of  the  eei'elielluin. 
Hasinir  liis  work  on  th<'^e  aMatuinie  ennelusions,  Van  Hijnlierk  lias  stiiilieil 
the  elTeet  iif  CirCIUnSCribed  extirpation  of  eertain  lulmles  nf  the  eereheliuni 
on  the  inUM-ulai'  fDnlriil  of  the  ililVei'ent  pails  ol'  the  limly,  with  the  rnlh'win"? 
results.  'I'dlal  ni'  par'tial  extirpalicm  of  the  loliulus  simplex  i>nHluees  side  to 
sidi'  oseilhitiiins  of  the  lieail.  indieat  iiiir  the  removal  of  t)ie  iuHueiieps  of  the 
eerehellum  that  eonti'ol  the  moveirienis  of  the  muscles  of  liie  rieek.  Complete 
extii'pation  of  the  ei'us  jirinuiin  of  the  luliuli  ansiformes  eauses  as  an  imme- 
diate irritative  elVeet  d\  Mamie  disturhanees  of  the  fore  limli  of  the  same 
side,  i-eplaeed  later  h,\  a  eondition  of  atonia.  whieli  makes  the  limh  liaufr 
limp,  and  of  asthenia,  whieh  makes  it  feelile  in  its  movement  when  it 
is  exeiteil  to  contraet.     I'Xt  irpat  ion  oT  tiie  i-ins  secundum  has  a  similar 


I 


I'  c  :l.  s,!,,!,.,  ,,i  rlu'  |.,nl-  "l"  '!;<  ":,mi'ii.ili.,n  . .  1 1  |.i  II  in;  ■.■■r.,i.|  •.iit  in  ."!■■  \-\.vt'  i  Al'li  r  r.ulk 
in  \,iu  Kijnli.  ik  ti..i-.  I. 111.;. II. i.  t'l..  .  II  I  On  tin  rmlu  >i.lv  ..i  On-  Imnrc  llii-  ril.itii.n  of  the 
.hlTtitiit  I.. I. till-.  1.1  flu-  I.:!uti.n.il  .i«  vt  |..|..i.«nt  '■!  lb'-  nui^r;ilii!!;rr  i-^  iTi.|u:tti 'I  ;uc<u.lnig  to  tlie 
III.,  ri     ..1     r...lU    i:..li.|    ill    Ih,     t<  M.       Ill, in;    I'aM.N'Ml    lilack.) 


intluence  on  the  ll^|sl•|.■^  of  the  hind  limh  of  the  corresponding^  side.  Extir- 
pation of  lioth  I'rnra  of  the  lolmlus  aiisifornds  causes  marked  asthenia  and 
atonia  in  lioth  fore  and  hind  limh  on  the  sjime  side  as  the  lesion.  A  eliar- 
aeteristic  distui'lianee  in  walkiiiL'  ilevelops  as  a  late  clfeei  of  this  extirpation. 
It  has  heen  termed  the  "■hen's  vmIi."  Kxtirpation  of  the  loliulus  i)ara- 
medianus  causes  rotation  on  the  longitudinal  axis  of  the  hody,  with  pleuro- 
thotonus  to  the  oper.ited  side,      i  l"iu:.  '2'J4.  i 

Just  as  in  the  ease  of  ci-rehral  loc-alization.  so  in  cerehellar  wc  find  that 
within  each  of  the  hir;.'est  centers  a  more  particular  localization  can  be  made 
out;  thus,  in  each  of  the  centers  lor  the  u|>per  and  lower  extremities, 
there  is  a  definite  ai-ranu;ement  of  sidisidiar\  centers  for  the  direetion  of 
the  activities  of  antajronistic  nniscio  v'roups  concerned  in  tlie  movements  of 
particidar  joint-.  It  mn>t  he  ri  Uieniiiered,  iiowevrr,  that  in  all  these  ca.ses 
no  real  paralysis  is  produced  hy  extirpation,  hut  only  a  want  «f  eoordina- 


i  ii 


H70 


•nil'.  <i  NTHxr,  MKVors  sy>ti.m 


,;.,„  on  a.-.-.„n.t  ..f  tl>,.  la.-t  llu.t  ll..  stl..-ni...  t..,.i..  ni.-l  stati-  in.p-.lsos  ir 
.luiml  for  inuscular  lianuoiiy  aro  not  properly  I'laboral.'.!. 

\ft.T  somo  tin..',  as  in  tlio  case  of  .M.inplcte  cerebellar  exIiriMtioi.,  tl.' 
sv.npton,-.  .M^ulualiy  .iisa,.p-ar,  l.ut  tliey  ean  W  ol.taino.l  more  or  less  el.ar 

u'eterisfi-ally  in  praetieally  all  anin.als,  at  least  in  ail  those  that  have  1 m 

investi-rated.  in<-hi«liii>r  ilo-fs  and  monkeys. 

I,  will  1 r  int.'rest  to  eonsi.ler  for  a  nicm.ent  tlie  possible  causes  tor  the 

„l,in,ate  .li-a, aranee  of  the  sy    pto„,s  of  eerehellar  eNt.r,.atM,n      'I  h-se 

are  either;  (1)  an  or-anie  eon.p.Misation  by  the  unin.jure.l  parts  ot  the  eere 
belluin    or  rii  a  fnnetional  eo.ni.ensation  by  the  v..luntary  eentcrs  ot  the 
eerebrnin      Althonizh  the  former  of  these  methods  of  comp-nsation   n.a.v 
so.netin.es  .hnvlop  after  partial  .lestruetion  of  the  cerebellar  cortex    it  ean 
„„t  of  coni^e  explain  the  r.covery  which  we  have  seen  to  occur  after  the 
entire  eerenellnm  has  been  remove,!.    The  most  important  compensation  no 
doubt  is  etTeeted  bv  the  eerebr.m.,  as  the  following  observat..m  clearly  n.^ 
dieates      If  half  of  the  cen.bellum  of  a  doj;  is  destroyed,  and  the  animal 
kept  alive  until  the  sym,.to.,,s   .f  cerebellar  extirpation  have  entirely  dis 
appeared,  it  will  then  be  found,  if  the  eerebral  eenter  on  the  opposite  side 
is  removed,  that  the  symptoms  return  in  their  ori-inal  seventy.    Alter  this 
.second  operation   the  powers  of  standinir  in  the  erect   position  and   ol 
walkin<;  are  perniai..^ntly  lost. 

CLINICAL  OBSERVATIONS 

Apidi.'ation  of  these  laboratory  results  has  been  reeently  made  in  the 
,.|inie  the  most  important  contribution  having  come  from  the  elmie  ol 
liaranv  who  for  his  work  was  awa/ded  the  Nobel  prize.  Tn  cases  of  abscess. 
cvsts  or  re.^i..nal  agenesia,  it  is  now  |.ossib1e  to  determine  the  exact  site  of  the 
lesion  ill  the  cerebellum.  To  effect  this  localization,  it  has  been  necessary  to 
work  out  certain  clinical  tests.  The  most  important  of  these  is  called  the 
■nnlr  r  ^  s7  This  is  described  by  Davidson  Black  as  follows :  ' '  The  patient  s 
eves  bein.'  closed,  he  is  asked  to  execute  a  simple  movement  m  a  given 

d'i, tion^with  one  of  his  o.remities.     For  example,   the   forearm  being 

firmlv  supported,  the  patient's  index  finger  is  extended  and  brought  into 
contact  with  that  of  the  observer;  tli.'  patient  is  then  required  to  move  bis 
fin-ror  verti.-allv  .lownward  and  then  to  return  it  to  its  previous  position. 
The  test  is  repeated  a  number  of  times,  both  in  the  vertical  and  in  the 
bori/onal  .lire.-tion.  and  if  any  tendeney  toward  deviation  from  the  plane  of 
movement  be  present,  its  direction  is  noted.  By  slight  modification  of  the 
foregoing  procedure  it  h  passible  to  test  eael,  of  the  limb  segments  in  all 
positions  of  rotation,  pronation  or  supination." 

The  index  lest  is  appli.'d   (V  without  previously  producing  nystagmus 


KINCTIUNS    OK    Tin:    CliKKHII.l.rM 


871 


and  (2)  after  produciiin  artifurial  iiystaKmiis.  Tlie  artificial  nvstagimis  is 
produco'l  li.v  siiiniiini;  tlic  i)a1i('iit  two  or  tlin-i'  limes,  aiul  eonsists  of  con- 
stant lateral  niovenionts  of  the  eyelialls,  ((iiiek  in  tlic  diri'etion  in  which  the 
artificial  nioveinent  took  jilaee,  and  slow  in  the  opposite  direction. 

In  a  normal  sulgeet,  previous  to  s])innin<r  tlie  index  test  shows  no  devia- 
tion, but  after  the  production  of  artificial  uystafrnius  a  deviation  is  noted 
in  the  directinn  corrcspoudini;  to  the  slow  jerk  of  the  uystagnuis  (reaction 


l-'iK^.  2J?  an<l  JJ'-  tti.u'^tnt  u  v|.<vinil>  tlic  iiil't  flati  tal  aii.l  ilu  pn^trrinr  aspect  of  ihe  Iitiman 
icrrlifllum  tii'liiatiiiK  icttaiii  rcrtlullar  I->i-a!i/aIH'iis  airMr.lcii,'  l-i  U.ii  I'ly.  (After  llaranv,  tr<ini 
Anclrr 'rhciiiiai  <t    Ihinipt.     Op,   cit)      N.    \ll,    N<rvii>    faiialis;    X.    I.\,   Xervus   G^|^^"l'l!aryll^,■t■^5; 

N.    XII,    XiTVllS    lis  I'(>yloS5U3. 

The  siKiis  111  ihf  ahove  itiauram  itKliiate  the  exact  Imali/atiuii  nf  llie  cctiters  for  the  tonus  of 
tht  inu^ciilaluit  ct.n.  criu-.l  in  some  <if  the  niovetnenls  of  the  rik'lit  arm  ami  liK.  t^  inarks  the 
center  fnr  ilitwuwaul  niuveinents  of  the  ami;  x ,  fnr  aluhiction  uf  the  arm;  O,  ahluctitm  of  the 
hand;  -J-  achl'i. 'liii  c.i  the  arm;  ^.  aMiicllm  of  the  hip.  X.  V.  hi.lic.itts  Xervus  triseminus; 
\.    VI.    Nerviis    a'^l'icins;    X.    \'M.    Xervus    facialis;    X.    IX.    Xervus    slo.^.ipharyngeus;    X.    XII. 

\'..r.....i       1.  \  .   .11.1....!..  I    |.'rti..i        I  )'i .' i  .1  vit.i        t'.lni'L'     ■ 


N'ervus    Ini.'.yh 


i  I-'rnni    I>aviiKnn     Clack 


deviation).  When  a  cerebellar  lesion  exists,  the  index  test  performed  on 
a  patient  without  nystafrmus  somotimos  causes  a  spontaneous  deviation  in 
the  sefrnient  of  the  body  oorrespondiiij:  to  the  positinn  of  the  lesion  on  the 
cerebellar  cortex,  but  more  frequently,  if  it  is  applied  after  the  production 


f  I 

f 


•  ji 

1 

r 

„  :;* 

hi 

't> 

;■< 

87: 


Tin.   ('KNTKAI.    NKUVOIS   SVSTF.M 


of  artilu-i..l  nvstajrnms.  the  nN.Hion  .Aviation  in  tl.at  sofrm.-nt  w.l    la.l    (. 
be  ol.tai.H-a.     Tl...  .-xa-.t  sif  of  Hu-  .M.n-lu.llar  lesion   is  .l.airn..s.Hl  ,mrtl.v 

from  tl.o  natnn.  and  .lirrrtion  of  ti,.>  .l.-viation   vvhi.-h   is  pnulu...,!  and 
,„,,,  ,•,,.„.  tlM.  M.vM„.-nt  ..f  tlH>  lin.l.  in  wlm-1.  it  orc-nrs.  the  ..x,.lana  ..... 

for  ti.o  .list...-l.an....s  luMnj.'  tl.at  inte.MVn.....  with  th.  ...-n-hoUar  ......t.-o    ..t 

.,„,.  n...s..U.  -.-o..,.  .-anM-s  th.^  antaironisti.  n...s.-..lar  !.M-.n.ps  to  rorfonn  th.'.r 
...ov.wnts  in  an  .■xair^r.Tat.,!  ma...,.-,  so  that  tl...  s..vM...«nt  ,n..v..s  too  n.n.-h 

ill  their  (lir.M'tinn. 

Maranv-s  .•oiwlnsions  so  far  n.ay  W  M.,i,n.ar./,.-.l  as  follows: 
(1)   Tl..>  .■.-..t.Ts   f..r  th..  .•xtr.-.niti.-s  a.v  l...-at.Ml   on   the  .-..H.-x  ..f  tl..> 
,,.M.,ispli.Mvs  in  th.-  s,.,..il,...ar  .snp..ri..r  an.l  inf-Tior^  an.l  .li-.-astri.-  lohulos 
IS,.,.  y\"  •'•'."<        The  ,■,.,. ivM.nlati....  is  une.M.ss...!  or  lioniolat.Tal.  thns  eon- 

trastin.^vith  .•.•.vl-ral  l.M.ali/ati.m.  in  whi.'h  it  is  ...•oss,.d  or  h.'torolat..ral. 

r>)  Vithi..  eaeh  ..f  1h.'so  ehi.'f  .•ent..rs  th.^ro  is  a  f.irth.T  loeal.zat...n. 

whi.-h  howov.-r  does  ,.ot  refer  t..  anat.m.i.-  ..M-....ps  ..f  ninseles  hut  rather  t.. 

,1,..  fnn..ti.mal  perf......a,iees  of  the  diff.-'ent  se.4...e..1s  ..f  the  l.ml.      Thus. 

uithi..   the  am.  .•...it..rs  there  a.v  subsidiary   ......ters  eone.-rned   in  the 

nioven.e.its  of  the  li,..!-  in  the  va.ions  ph.....s  i..  n.tati..n,  m  pr.mat.on 

a.i.l  in  snpi.iati.....     It  is  a  f....eti,...al  rather  than  a..  a..atonue  loeal.zat.on. 

(•!,   Wh.M.  a  e....ter  eon.M'rn.-.1  in  the  ni.nvn.ents  of  the  limb  in  a  certain 

direetio...  ...!.'..  to  the  rijrht,  is  suddenly  d.>stroy.>d,  a  sp..ntan.-..ns  .h>v.a- 

tiuM  is  pro.l...-..d  in  the  opposite  direeti.m   fto  the  left). 


(  IIAI'TKW  (• 

TIIK  (KI.'Kr.MI.I.I  M  AM)  Till:  SK.M  K  I  i;r||,Ai;  (ANAI.S; 
KINCTIMNAI,  TKSTS 

'I'lii'  (•(■I't'lM'limii  sci'M's  iis  Ilif  ^.TCiit  iii'ivi'  ciiiliM'  In  wliifli  life  tr:iii>iiiit- 
ti'il.  tliniiiL'li  llic  Viiiiiiiis  |ifo|)ri()c('|it(ii's,  tin'  inipulM's  wliicli,  iis  it  wen', 
iiil'itriii  it  ii>.  to  tlif  I'XiicI  ili'L'ii'i'  III'  iiHioiMilai'  clVui't  I'l'iniircij  tu  iiiiiintiiin 
the  .-iiiiiiinl  ill  its  \;niiiiis  |mis1  iii;s.  It  is,  as  Slicii'iiiirtciM  puts  it,  tlic  iicaij 
iraliiilinii  lit'  till'  pidplinccptix  I'  systilii.  Sllrli  iiiipilUis  t'lniil  tile  llllisclcs, 
ti'iiiliiiis.  I'ti'.,  ciiiild  111(1.  iiiiwcvii'.  supply  iiiliiriiiatiiiii  icL'aiilin^  tlio  cxai't 
pKsiliciii  (if  ihc  liody  in  space.  Fur  this  purpose  special  receptors  ciiii- 
Mected  with  the  ei;;htli  nerve  ale  proviiled  in  the  si  mii  iroihir  fiiiiiilx. 
These,  it  w  ill  lie  reiiieiidiered.  arc  three  in  nuiiilicr  on  either  side,  each  canal 
consistintr  of  a  semicircular  hone  liilic  attai-heil  to  the  vcstiluilc  of  the 
internal  car;  and  th-y  are  arranired  so  that  they  lie  at  riirht  anjrics  to  one 
another  in  the  three  )>laTU"s  of  space.  The  three  canals  on  litlicr  side  are 
thus  disposed  so  as  to  form  an  arrangement  liki>  a  \'  sjiaped  armchair  with 
the  liai'k  inwards.  This  arraiie'eiiient  causes  the  posterior  vertical  canal 
cd'  oni'  side  to  he  in  the  same  plaiii'  as  the  superior  MTticid  canal  of  the 
opposite  side,  the  external  canals  hoin;.'  in  the  liori/ontal  phinc  on  hotii 
sides.     The  aiauyeincnt   \mII  he  plain    from   the  diaL'iaiu      l-'iir.  1227'. 

Within  the  ossemis  canaN  are  suspended  iiiemliianons  tnlies.  which  do  not 
(ill  tho  canals.  The  canals,  etc.,  contain  llnid.  hut  are  not  completely 
tilled.  The  osseous  as  well  as  the  memliranous  canals  arc>  dilated  at  oiw 
end  to  form  the  ampidla.  and  it  is  here  that  the  vestilmlar  division  of  the 
ei^'lifli  nerve  ends  in  a  structure  called  tl'e  "iTista  aeustica."'  consistine-  of 
hair  cells  supported  hy  sustentacular  cell-.  The  ner\c  terminates  in  tine 
arliori/alions  lielween  tin-  hair  cells.  In  the  s.iccuie  and  utricle  mi'c  strne 
lures  similar  to  the  crista,  e.-jljed  the  iiMi-uhe  aeuslii-;e.     Tli(vi>  stfiicluies  aii' 

n ptors   specialized    for   Ihc    purpose  cd"   respondiiiL'   to   diaiM.'cs    in    ihe 

position  id'  flie  liead,  and  thcreforo  of  llio  hody  in  u'cneral.  When  the 
head  moves  in  n  certain  jilane  of  space,  tlio  tinid  in  the  memliranous  canals 
and  in  the  utricle  and  saccule  on  account  of  inei'tia  underv'oi's  a  certain 
movement,  wliieli  acts  on  the  hairs  of  the  hair  cells  and  thus  sets  up  a 
stimtdus.  Accordiii)'  to  the  de>_'roe  of  the  siimnlation  in  the  various 
ampulla',    which    a<;ain    will    he   dependent    upon    the    plane   or    planes    in 


i  Ml 


which   the  movement   of  tl  e  head   1 


las  oecnrred.   impulses  are  transmitti 


s;:! 


■.9.i:.T    1* 
'    1      I  ' 


:l|!.'!i 


Hf  if;! 


r  is  1  •! 


!'■' 

.  1 

1 1 

1 

! 

1 

' 

' 

<  ■ 

1 

■' 

'  ill 

874 


THE   CKXTKAI.    NKKVOIS   SVSTKM 


tlm.u^'h   the  v..stibul..r  imtv,   an.l   tl.csr   in.imlsos  ultimut.-ly   ri-udi   th. 

The  cxiHTiiiu-ntal  .•vi.l.iu-.-  for  i1..m-  ,.n,u-lus..ms  rvii>m\mK  tl..-  fuiictu.n> 
„f  tl...  scniuMn-ular  n.nals  is  vory  stn.n^-.     Tims,  aftor  .U-strm-tum  of  all 
th..  ,.anals-an  ..,u-rati..n  whi.-h   is  ,mrtioularly  .asy   in  the   ,,.^^■o..^-th,. 
„,.i,„al  behaves  very  n.ueh  the  same  as  after  eerehellar  .h^st  met ......     Aft.-.- 

so,..e  n.,mths  th.se  ,lis..nl..rs  .lisappear.  h.eause  the  ..en-lM-ll,.,,.   l.-an.s  1.. 
...mtrol  the  .....ve....'..ts  ..f  th,.  ho.ly  f......  oth.-r  pr..pri..ee,.t.v.-  ....pulses. 

nartie..larlv  th..se  of  si^'ht.     If  M.eh  a  .•.-.•..ve.v.l  a......al  .s  ha...h.p..l.  th.- 

'v,..pt..n.s  ".vt......   i..  all  severity.     This  ...,...p.'..sati....   is   f„rlhe.-......v  an 

ea.ieative  p.-.-e.-ss,  for  it  .h.es  not  oeenr  wh.-..  the  e.n-l.ral  .Tnters  as  w.-ll 


KiK 


ml.ir 


,aK   ,.l    til.-    .a.,   showinii   f.i.-.r    arranKn:i.-..t    ■"    Hu-   thrfi-    planes    ot 
•  laiT.      <Fr.im   .luwcU's   I'hysiclogy.^ 


,us  the  s..,..i.ire..lar  eanals  are  renu.ve.l,  a...l  it  ean  he  abolished  in  a  re 
eovere.1  anin.al  bv  rc.ioval  of  the  cerebral  cortex. 

Manv  ..bservations  of  trreat  int.-rest  have  been  lua.le  .•..neern.nj;  these 
labvrinthine  s,.nsatio„s  by  I'ike  a...l  ..th.rs,  bnt  we  ea..  not  .l.seuss  th.-m 
further  h.-r.'  <»ne  point  ..f  interest,  h.iwever.  is  that  fore.-.l  movements 
in  definite  planes  are  i..<h.ee.l  by  .emox;  1  of  a  eanal.  Removal  of  the 
hori/.....tal  ea..als,  f.,r  example,  eanses  e..nti....e.l  noddinp  m<.veme..ts  «t 
the  h.a.l  in  the  plan.-  of  the  i...jur.d  .-anals.  An  exper.me..t  of  srr.-at  s.-- 
.lifi.-anee  was  p.-rform.^.l  by  Kwald  to  show  the  eiT.-et  of  eaus.n-  a  move- 
ment of  the  llui.l  in  one  ..f  the  ea..als.  For  this  purpose  a  bony  eanal  was 
ope..ed  at  tw..  |.l.. -es  bv  a  dental  drill.  ThroUfrh  the  hole  farther  f.-om  the 
■uniniUa  aina!-ri.i  •.■.■,:-^  iMtr«due.>d  so  as  to  block  the  backward  movement 
of  fluid'  and  Tnto  that  n.-arer  the  ampulla  a  fine  tube  was  i.iserted  eon- 
,.oet.Ml  with  a  rnblur  bulb.     Uy  manip.ilatinsf  the  bulb,  the  membrano..s 


ciuiiii  i.r.iM  AND  Tin:  si:MinR<i  i.ar  ianai.s 


87.-) 


Piitiul  I'oiild  lie  t'Diiiprt'ssi'd  anil  ciirrciits  set  up  in  the  ciitlolx  inpli.  It  was 
fouiul  thut  till-so  ciirriMit.s  iilways  i-aiisod  tin*  uninial  to  nuivi-  its  head  and 
ryes  in  tlio  plani'  of  tlio  I'unal  that  was  lipinjt  stiiniilatrd  and  in  tlif>  diri'i'tioii 
(if  till-  iMirri'iit  of  ondolynipii. 

Finally,  visual  inipri'ssions  supply  niui'li  of  tin-  infornuition  tin-  imti'Im'!- 
Iiini  roiiuiri's.  tin-  cliisi-  assoriation  iit'  tlir  iiiovi'iiii'nts  of  tlio  I'Vi'lialls  with 
I'pri'lii'llar  and  laliyrinthint'  disturhanrcs  liiMn>r  wi'lj  mm  oirnizi'd.  Thi'  nys- 
ta^inus  already  dcsiM-ilii'd  in  conni'i-ticin  with  Mavnuy's  tests  is  dependent 
upon  this  assoi'iatiori.  The  symptoms  and  seiisatiims  iit"  jiiddiness  or  iniusea 
produced  liy  rotation  of  the  hoily,  or  hy  uiuisual  movements  sueh  as  those 
of  a  lioat,  are  no  iloidit.  due  to  the  irrejiuiar  and  unusual  variety  of  laby- 
rinthine sensations  whieh  they  exeite. 

In  n  word,  then,  the  f\inetion  of  the  eerelielluin  is  to  reeeive  proprio- 
eentive  impulses  from  the  Imdy  alou};  with  lahyrinthine  and  vistinl  ini- 
])ressions  and  \i,  intejrrate  and  develop  from  them  impulses  whieh,  h.v 
lieiii).'  transmitted  to  the  eerehral  and  other  nerve  centers  that  dominate 
museular  movements,  so  coordinate  the  nerve  iliscliari;es  from  them  that, 
when  muscular  movement  occurs,  it  does  so  in  relationship  to  the  previiuis 
position  of  the  aninud  and  in  the  most  elVicicnt  way  to  attain  the  object 
for  which  the  movement  was  made.  The  cci'cbellum  is  the  head  nucleus 
of  the  jiroprioceptive  system. 


THE  ASSOCIATION  BETWEEN  THE  EYE  MOVEMENTS  AND  THE 
SEMICIRCULAR  CANALS 

The  elo.se  as.sociation  between  the  eye  movements  and  the  seniicircubu" 
canals  is  indicated  by  the  occurrence  of  nystafrmus  when  the  car  is  .stimu- 
lated either  electrically  or  by  means  of  moderately  coM  water  impinginir 
on  the  membrana  tympani.  The  latter  method  of  inducin<r  nystasjmus  is 
styled  the  caloric,  and  is  employed  in  the  examination  of  candidates  for 
the  aviation  service.  Its  value  over  the  tests  of  nystajrmus  after  rotatiiif; 
the  body  and  the  index  test  already  described  depends  on  the  fact 
that  it  enables  us  to  test  each  vestibular  apparatus  sci)arately. 

Water  at  (iS  V.  is  allowed  to  I'uii  throuirh  a  stop  nozzle  into  the  ex- 
ternal avulifory  canal,  free  of  wa.x,  from  an  irrifratioii  bottle  placed 
about  '-i  feet  above  the  head,  which  is  meanwhile  tilted  at  an  an^le  of  30^ 
forward.  In  about  40  seconds  a  rotary  nystairnius  with  the  direction  of 
the  jerk  to  the  side  o])posite  to  the  douched  ear  should  be  evident,  or 
dizziness  complained  of.  The  reaction  test  is  then  applied  and  im- 
mediately afterward  the  head  is  inclined  at  an  aufrle  of  fiO''  backward 
from  the  perpendicular,  when  a  horirontal  nystagmus  to  the  side  op- 
posite to   the   doiu'heil   ear  should   develop.      Ueviation    is   ai:ain    tested. 


5  1  ' 


,p;        ,1 


I       : 


;iiii-i 


s7t; 


III     I  I  Mini     M  IIVHI  -   .■.V>TI.M 


,,.,,,,„.,.,,,..,.,..,.. n,...!  ..M  .1,..  ...IMT  ...r.      int   lak.s  l..n«.r  llw-n    .. 

.,. Is  f..r  tlu.  .n.t.umus  I..  a|.|MM.,  i1m>  v.stil.ul.r  aiMmiMlus  ..l   tl,.l 

s,a..  is  .■.l.,M.>n.;,l.    Al.s.n I'  Hi."  .-..•ti U■^u^^u>u  .(U-v  11 ....-InM..'  .^ 

i,  (■."ttiiiii  >iyii  "I   iiH. mill  .Ml'  (lis.'ii>'v 

1,    is  MH.InuM.-llv  .ssmtinl   llial   tlirsr  t.sis  sli-ul,!   Im.   n.nst    ..aivln  Iv 
.p,,li...l  t,.  all  unni.l  !..■  aviiitnis.     Tlu.y  !r. ntly  .wval  I^Mons  ol  ll.^ 

,l,.,„s..h,.s  iHTlVcMv   Monnal.  an.l   uh"  in.l.v.l    may   lias.-  l-oaM.'.l   -I   th.'ir 


,„.„,.rsor...,uitil.nn."  iM-au- tl-.y  in.a«i.u..l  Mial  Ir Lmihuiu  soasi.-K 

',„.ss  ,„•  .-ailun-  to  I... ■  -li-    ii.  'i-'"'"^'   ""'"-"-'  ■[  '"«"  ;'-•'"'••""" 

,„•  this  lunwion.     Th.-v  .-an  1-  l-.«.i'»  t''^"  •"='">  ^'^ '"";-  ''"^"  '''•""■  '" 

,l,..ir  .l.alh  !K...au.-  of  impair nt  i>,  th.  .ar  n>...l imm.      NMm'm  o„      t.-rr, 

,i,,„a"    ,1...   muMM.lar   s.-ns.-   a.i.l    .•nlan.ous   s..usa1.o„s   ot  rn    ...ak..   11.. 

v..stil,ular  waUniss  of 0.................  Mi.   .h-n  .l.,..  iw.l  o,  ,1,..^.  n.n 

1,i„uto,v  s.<nsat,oMs  a.Ml  .l.MM.n.lml  on  tlu'  ..ar-lmla,,,...  nu.,-l,an,s.n  alon- 
as  in  lIxiM..',  any  u.aknrss  1m 1.1-  a  s.-iious  ux-uiu-v. 


i  % 


m 


lis    i  ■« 

Ijj.i  i 


(■IIAI'TKU  ri 

Till-;    AITONuMIC  XKKVOIS  sYSIKM 

III   •li>iii>siinr  till-   plixM   lui;\    1)1"  ilic  central   nci'Vons  Msti'iii.   we   Iuim- 
lijokiri   ii\Miv   frniii   till'  tia.iilii)iiiil   ti'xiliouk   tn'.itiiu'iit   of  the  miIiJ.mM    in 

lii.ll    \\i-  I    MM'  !(  11   IHMi'liciilly  IllitiiUi'lii'il  ;ipiy  (lcM-n|itii I'  tin-  iMillIsi-  iif  tllf 

Viii'ioiis  iMTVi'  inii'ts  111-  ol'  tilt'  iiositiiiii  (if  flic  iicfvi-  i-i'iitcrs.  Wi'  liavo 
piirsiicil  this  imlicv  in  the  lii'licl'  tlial  tlii'  study  of  tliosn  details  (if  struct  lire 
liclipii!;s  Jiivt  as  surcl.v  to  tlic  analoiiiist  as  ddcs  tlic  structure  (if  (itlicr  i)nrts 
(if  tlie  ImkIv,  iKitw,  .  staiidiiii;  tluit  In  trace  tlie  course  of  tlie  nervous  patli- 
ways  lie  iiia,\  have  to  call  tii  his  aid  llie  pliysiolojiist  and  clinical  iieiirolojjist. 
There  !•.  line  part  of  the  nervous  system,  however     iianiily.  the  involuntary 

"I"  autoliiil the   ph    NJoloiiy   of  which    it    is   iiupossililc  to  discuss  apart 

I  idiu  its  anatiiuiy.  Iiccansc  this  has  depended  very  larirely  on  pliysiolojric 
methods  ('ly  its  elucidation.  I'litil  siicli  nietliods  wire  iinphasi/ed  and 
rvhiie  anatimiy  alone  was  de)ienili'd  upmi.  little  could  he  learned  of  the 
functions  and  connections  of  the  sympathetic  chain  and  of  the  various 
nerve  plexus  tliat  compose  the  involuiilary  nervous  system.  We  bhall  liere 
revi(nv  lirictly  the  '_'eneral  anatomic  plan  of  this  system  at  ileseribed  liy 
Caskell.'' 


GENERAL  PLAN  OP  CONSTRUCTION 

Tlie  plan  of  the  involuiilary  iiervens  system  is  much  the  same  as  ihat 
i>r  the  Miiuiitaiy,  the  main  ]ioints  of  dilTerenee  Iieiii!;  dependent  upon  the 
liicalion  of  the  neliroMs  coiiiposiiifi  the  icllcv  arcs.  It  will  lie  rememhered 
that  there  are  tliieo  of  these:  the  receptor,  the  internuncial.  and  the  .f- 
iictiir  I  iinris  I  pa^re  "S'J  i .  The  icccptur  iiciiinns  have  the  same  posit  •\ 
t'urlioth  systems;  namely,  the  posterior  root  ^'anjilia  i  Kij.'-  -"JS  > .    The  iiiter- 

iii'iicial   neiii s  of  the  iiivoliintacy  system,   like  ihoso  of  the   voluntary. 

have  theii-  cells  ill  the  spinal  cord,  where  tlicy  ,ire  represented,  in  the 
llinracic  ic'.'inii,  1,\  the  cells  of  the  lateral  horn  «{  ^.rpay  matt  •■;  in  the 
sicral  rc'^'ion,  \<\  a  similarly  placed  collection  of  ells;  ,iiid  in  the  l-ulhar 
ic-ri'in.  mainly  liy  the  dorsal  nucleus  of  the  vai;ns.  The  main  cause  for 
the  (litTerence  hetween  the  two  systems  iN  dependent  on  the  course  of  the 
fillers  (if  the  iiiti-n:i;niial  neurons,  in  the  involuntary  system  fla-y  leave  the 
spinal  cord  In  fun  coniu'ctiiipf  vith  the  ofToetor  neuron  nerve  cells,  \  hieli 
aii^  ei'iitained  in  the  various  «;ain;lia  found  tlii(nis.'lii>ut  the  body,  ^\llereas 

877 


i    IV 


«t 


H7H 


Tin:  I  f  NTK.vr,  mhvois  kvhtkm 


r  njii-;; 


l5f:r 


#;  ' 


i'l 


npi 


i-ti 


I 


i  I 


:H 


ill 


y 


ill  the  volimtiiry,  they  reiniiiii  within  tht-  spinal  cDnl.  hikI  terminate  on 

the  flTt'rttir  iii'urnns,  wliirh  iiro  thr  nnt('tii>r  hnrii  fflls. 

Tlio  outtlow  t'roni  the  spinal  <-iini  nl'  iiivoluntnry  internuncial  (iherx, 
which  we  sliali  hiTcaftcr  >\\\v  ciiiiin ilor  fillers,  nconrn  itlunK  the  an- 
terior spinal  roots,  hut  is  somewhat  iire^nlar  in  ilistrihiitinn,  heeanse 
it  is  iiiteiriipted  in  tiie  eervii-al  ami  liinihar  retrions,  where  the  nerve 
]>iexus   for   till'   extremities  eonie   in.     There  are,   therefon-,   three   main 

regions  of  outllow  for  tin mi;'etor  lihers     a  thorueieolumhar,  a  hnlhar, 

and  a  sa<-ial:  anil  tlie  tihers  (  Kiif.  'J'JM i  tlo  not  liehave  in  the  saine  nwinner 
in  all  of  them.  Tiu'  fibers  of  llie  thoraeieoiumhar  r-<'t;ion  form  tlie  so- 
ealleil  siiniinitln  lir  siistiiii.  anil  nni  l>y  tlie  eorrespoiiilinij  vihite  rami 
eomiiiunieanteH   either    imiiiei|iatel,\    to   the    uanitlia    of   the    synipnthetie 


I-'iK  J-'."*  1 't.inr.tm  •«.  ilhi'-tr.Uf  tl  i-  ttirtrrrtil  aTr.u'Kiiiu  in-*  itf  llw  inti-i  iniiu-u1  nriironn  "I  the 
v.'hinlary  aiiil  invotnnlary  turv.ni-*  sj^i.nT*.  In  liotli  f^x-^lfiiw  tin-  afffitiit  hht-r  ttrminatrs  (liy  col- 
l.-itrraNl  aroiiii'l  a  t  rll  .if  tin-  ki  ay  ni.ittir  nf  thn  ctmi.  In  thr  Mihiiitiry  .ly-.tfrn  tin*  cell  t!i  Slt- 
iiati'-I  III  till-  pM^tt  riMT  liMfn.  .tii.I  its  ax'in  iravrU  to  an  antrri'ir  hnrii  n-ll  In  thr  mvnliituarv 
«y*trin,  on  the  fthir  haiiil.  it  i^  JMtati-il  in  thi-  I.ltfr:il  h-irn,  ami  iIh  axon  leaves  the  cnril  liy  the 
anteri'ir  nmt  ami  IraviK  liv  the  white  rannis  into  a  xymi-athrlu  KanKtmn,  where  it  connect!!  with 
a   nerve    rell,    w  hi.^i     ,i\i,ii    form-,    the    intstKaiiKli-'im     tiher.       (  l-'nini    ('.a*l<el].» 

ehaiii,  <ir  hv  the  splanehnie  nerves  to  the  aliiliuiiinal  K>i>i^lhi.  In  the  Kan- 
uliii  are  situated  the  i-ells  of  the  efTeetor  neurons.  The  fibers  of  the  saeral 
reifion  eonnect  with  efTeetor  neurons,  forniiiii;  the  pririr  finuijlionic  (irniif) 
(pelvie  nerves,  iiervi  erijientes^  ;  and  those  of  the  bulbar  outflow  with 
efTeetor  neurons  loeateil  either  jieripherallv  or  in  the  <ran(;lia  of  the 
vaf/iis  ami  the  seventh,  ninth,  and  eleventh  eraiiial  nerves.  In  the  mid- 
brain there  is  a  fouith  irrinip  of  involuntary  eonneetor  fibers  runninR  to 
efTeetor  nenrinis  found  in  the  eiliary  (laiiiilia. 

The  anterior  roots  of  many   of  the   spinal   nerves  are   therefore   not 


"     #7  • »' 


\: 


!i 


lii; 


-.il,lt      |1.   X^ 

.  !■■... til    11. 


IlL.J. 


).  1 1..- 


II.       -\  '.  I  .til'.  -1        M.      \    •■        ^  -f.  ■■       I..     '>i        ix.l      'I.. I 

I     1,.|.      ,1,  I     1 -If"cl I-.1  A.        X       .     

..■1^,m:      /       -i.!:,,:,     1-..  ,i:li.  II.     \  I        iri.  u     -r'!-""  I"i" 

t r1.   :..       Cl'iulu;      -I         l.M  ..U.I^tU.       I!it\<v.       \     / 

II 1      ■,.n.       ,..,  .l.-.li..  r'l.       M-...„.       ,      ,.,  ; 


11        Till 

ll      ll,l"^I...M 

tr.cll,  ..1 

l:.  I.        111. 

M..I         IN't.. 


i      !  !  I 


m  :fj|j 


.  1  k 


^ 


n 


>v.\ 


1 


-f-: 


r 


y^dX^': 


-Jpinal  cord 


.:   -   .Svmpathetic  chain 


'';->Jolar  ganglion 


'■'"         ■■''         l''''t ill.i       l.mtl-li      -l! ni;     ll„      ,n, i      ,,,,,„,-.,,.„,      ,.,      ,|„       ,| 

inu     llu      K1...I     N'l. nil,      ,„r»,  Th.      I,  n   h.,n  I     . I, .,„,.„,,      :,,,,,,.  ,,>-     •!.,       ,,--,,1      ,.       Mi:,  im,  ,,l     'll„ 

p., u-iMw  .......     :,:,,-    ,■.1,.:,,    ,„„,„.    ,i,.,„,,i     ,i,     ,.,.,,,, -in,.,ti,.n ,.„    .,,„|    i.',^,,,,, 

.1...,        ..-,  ...,1...,„       ,„       ,|„         .,:|..,^,.,J,     „„        ,,..„,        „1.,.|,        ,|„  ,....U..,„^I,..„„  ,,|„,-        ,,..1,        ,1, „,,V. 

'■■  ''"  '•■  'I""   ■i.-i,.,.,i.,.>.  .,1.  .,t  .>.,    1.1 1  >,.„!,,     ri„    ,,„|„| 1,,^,,.,,  ,,„,„,    ,   .., 

I  M  .  i.t.'.i..il     a,  ...T.i;.  ....  ..v 


...!.|.. 


il.il 


in 


ii  iil!  1 

I      ! 


N  I"- 


I  ! 

i    i 


:il     11 


THi;    .\l  TIINOMIC    XKKVOI  S   SYSTKM 


87!» 


<'onip<»sc'fl  piitirplv  of  filxTs  l>ol<>ii!.'iii(i'  to  volmitHry  rlTt'cfui'  neurons,  but 
also  of  conrioofor  (iliors  of  the  involuntnry  system.  Tliosc  aro  distin- 
trnishnl  iioni  lli4'  voliintMry  filn'is  liy  Ix'iinf  iiiucli  siuiiUcr  in  diameter; 
indeed  it  was  liy  this  cliarai'teristic  flint  (Jiiskell  sin'ceeded  in  traeiiiir  the 
course  of  t''c  in\  iiluntai'y  tilieis. 

In  liiii't',  thcr'fori'.  the  tilicis  of  the  iMtcrniineial  neurons  of  the  volun- 
tary iieiNoiis  systi  (ii  are  confined  within  the  I'ciitral  nerv(Mls  system, 
when"  tlu-y  are  contained  mainly  in  the  while  eolmnns  of  the  spinal  eord,  the 
)).\ramidMl  trai'ts,  for  (xam])!*',  Iieiiii;  eomposeil  nf  internuneial  fihers 
from  the  c'crehral  neurons:  the  eorresiMiiMlini;  tihers  of  the  involuntary 
nervous  system  iconneetori.  rui  the  dthei'  hand,  inn  out  cif  the  e.-ird  with 
the  anti'i'ior  roots  to  efTeetor  nenrous  siiuateil  cither  in  the  ({aimlin  of 
the  sympathetic  chain  im^  in  peripheral  |oealiti<'s.  .lust  as  the  voluntary 
internuneial  (iliers  j{i\  ••  o(T  many  <'o|laterals,  so  do  those  of  tlie  involuii- 
tai'v  system,  so  that  an  imjiulsi'  traii-niitled  liy  one  internuneial  neuron 
ma\  excite  a  liroad  field  of  effectors  We  shall  see  later  tliat  it  is  tliroujjli 
these  eollati'rals  that  reflex  responses  ciin  apparently  often  lie  I'xeited 
liy  the  stimulation  of  the  central  ends  of  nerves  such  as  the  hypoirastrie 
to  the  liladder  after  all  connections  with  the  central  nervous  system 
have  been  severed.       Fijr.  "ilJO.  i 

To  elucidate  Ihi  fiirllnr  iniirsi  of  the  iii\  rihinlary  fillers,  ainl  deter- 
mine tlie  location  of  the  elTector  neuron  nervt IN,  it  becomes  necessj\ry 

to  supplement  anatomic  with  |ih.\siol(i":ic  methods  of  invesliiration.  The 
\aiious  functions  of  the  innervated  parts  \asiMilar  chaiiLies,  muscular 
mo\enients,  irlandular  aeti\ity  are  observed  by  the  usual  methods  of 
the  phv  sioloi;ist.  and  the  ni'l'Vc  roots  or  nerves  believed  to  contain  the 
involuntarv  tibeis  either  cut  or  stimulated.  It'  a  chanire  is  obscrveii  in 
the  functions,  it  indicates  that  part  at  least  of  the  involuntary  nerve 
supply  is  contained  in  the  lu'i've  siiiiclure  that  has  been  cnt  oi'  stiiiiu- 
lafed.  Snidi  a  result  does  not,  hovvevi''-,  inform  ns  ,is  to  whether  the 
fiber  is  that  of  the  conm  (  tor  or  elTector  nenron  v  hether  it  is  ju'e 
■.'anirlionic   or   postv'aii'rlionic.      This   mav    he   determined    in    many    eases 

by  observinjr   whether  nerve  ili'Sienerat  ion   'Urs  as   a    result    of  iiittins: 

the  fibers,  hut  the  most  useful  method  foi-  ansvveiint:  the  (|Ucstion  is 
that  discovered  by  l.anirlev  by  the  use  of  nii-otine.  which  in  certain  i-on- 
ecntratioiis  specitically  paral\/cs  the  synajitic  connections  between  the 
coiHieetoi-  and  the  efVector  neurons.  If  a  weak  il  jier  ci'iil  i  solution  of 
this   .ilUaloid    is   painted   on    a    uaiijrlion    or   peripheral    uiMve    plexus   in 

\vliicli   tin nnector   neuron    finds   its   elTei-tor   ncr\e-cell,    it    will    break 

the   ni'ivi-   path,  so  that    jiliv  sioloifje   responsis   jiroduceil    l.y   stimulatiiiLr 
the  |ii'e','an!.'lionie  fibers  beconu'  no  lonjier  elicit.'dile      When  the  involun 
tary  conneelor  libeis  run  lhrou<;h  several  '-'antilia,  as  in  the  svnipathetic 


^(1 


IMI     ilNIKM.    MKVols    >Y>II,M 


h.    1 


S-        1      h 


!       » 


i  :;  « 


^   .       I 


•liiiin.  ii    liii'.iiiHs   iKisNililr.  Iiy  s_\>.tcm;ilii-;illy   iiiiimiii!,'   the   u'Jiiiiiliii   willi 

ilicotilK  .     til    till     r\;irll\      ill     wliii'li    111'    lllt'IU     the    IiImt    lillds    its    clTcclDr 

iiiTve  I'l'll 

Tilt'  ciMii  sc  .'I  I II I  I'liiicl  imiv  111'  I  111'  riVi'i'tiir  lirliriilis  uf  I  he  111  let'  nil  t  flows- - 
liiilliMr.  lliui  arlrnliiiiiliiir,  .■iiiil  s;icr;il      \.iiy   in  iiiiiiiv  (ji'tails  iiml   iiilist  In- 

I'.ilisiilili'il    sr|i;i;;|tl'!>-. 

THE  THORACICOLUMBAR  OUTFLOW,  OR  SYMPATHETIC 
SYSTEM  PROPER 

'I'lic    i-Hliliri-lur    tillers    ;ili-    -.jiiiiplv     I'lilililiiii     ill    tlicir    lilltflnw      iViilll     till' 

i-uril  lii'twiiii  the  (ii>l  lliiii;ii-ii'  .•iinl  tlic  ruiiitli  liMiilini'  sci_'iii('iits,  niiil 
lliry  iiin  li\  till'  wliiti'  iMini  rniMnniiiicjnilcs  tn  the  s\  iii|i;illicti('  i-liiiin, 
w  licic  SI  111  II  ■  111'  tlii'iii  i'iiiiiiri-l  w  itii  I'lTi'ftiir  iii'i'\  r  I'clls  in  its  yrnnirlia,  w  liili' 
■  itlicis  inn  liiMiiiiI  till'  rliaiii  tn  liinl  tlii'ir  rlTi'i'tnr  i-clls  in  I'lillati'ial  ■.mii- 
'jlia  ic-|irr'M  iitnl  li\  till'  sciMiliinai-,  sii|i('iiiir  aii<l  infiTinr  iiirsciitrrii-  ami 
till'    renal    in    tlie   aliilniiieii        'I'lie    lilurs    nf   tlie   efTeetm'   eells,   (ifleil    calleil 

)n>Nti.'aiii.'Iiiii  i''.   are   iji^t  inmiislnil    fimn    the   eiiiineetnf   nr   |iri"_'aiii;liiiiiif 

(iliel's   Iiy    liiiii'j-   liiilillleiliillaleil.      'I'lluse   ileliveil    fliilM    eells    ill    the    latiM'ill 

■~\  iii|iallietii-   !.Mii<r|ia    |ii- eij   to  llieir  .|i  vtiii;iiii,|i   eitliei-  Iiy    way   nf  iho 

•Jia\  laiiii  i-'i|iiliii|liie,iiileN  tu  till'  ^lyiiieiila!  liel'M's  al'lir  llie  fusion  nl" 
llie  aiileii.ii  aiiil  |insti'iiii!-  >|'iiial  rniit^.  ill-  Iiy  the  miter  walk  nt'  tlie  lilimil 
N.-.U         j-i'j.  'J:!l  ' 

'I'll.'    elVi-.-tni-    heiiiiihN   vriiiiJN     tlie    fiillnw  ill'.'   ^t  niet  llles: 

I  Tlie  I.!. mil    \esvels  anil  lieart. 

■_'  Till     iiiM^eiilatnie    nl"   tlie    sweat    ijlainls. 

"•  Tin     liiilselllatnie  n|'  llle   liair   t'l  liieles   allil   ntlier   liinseli'-,   l\iinr   iimli'i' 

!!  1  -kin. 

I  '!'lle  tllll-ellhllllle  nf  tlie  sn-ealleil  -i<.'ll|i  lit  a  I  illlel,  wllii'li  is  |-e|ire- 
-eliteil  ill  rlie  ailllh    li>    tlie  iiteiiis.  I''alln|iian   liilies.  ureters,  ete. 

."i     Tl,e    -liiiiiieters    iif    tlie    intestine 

lie'_Mri|il|i_'  the  innervatinn  nf  the  lilnml  r<<\. /\.  the  e\aet  sitllatinil  nf 
the   '_'aii';lia   in   \' liieh  the  etl'eeti.r   neiiriiiis  are  -itnatiil   anil   nf  tlic   iiel'M' 

Mm!-,  whieh  eiiiitain  tin niieetiir  lihers.  is  slinv\n    III  the  aei-niiipaiiv  iin.' 

t.il.l-       |.aije  ^>-l 

It  is  eleai'  thai  the  inner\  at  inn  nf  the  lilmiil  vismU  i>  |iraet  ieally  emi- 
lililli.lis,  the  elTiet.,1'  heljinlis  heinir  situate.}  Iinlh  III  the  lateral  alnl  ill  tin- 
enUateral  ehaiii  i<\'  '.'aliulia  Tlmse  nf  the  fm  iiier  run  tn  the  \essels  nf 
-1 1  iirt  lire-  iiiiK  r\  ateil  liy  the  eranial  aiiij  s|iiii.il  -e'^ineiital  iiir\e>.  while 
tliiiMni   ill,    lattei    sU|.|ii'.    the  M-seU  nf  the  aliilninilial  ami   |iel\  ir  \  iseera. 

The  e,.iile  elnr  lihils  In  the  <ll-,,ll  ,//.(in/\  are  aKn  sirietiv  enlililleil  In 
the    i  li.il  .■nlenlllllllia  1     -\-.leln.    the    eell     -latinn     liellrj     t.M|||,|     in    iji.      u.ill'jlinll 


ill  n 


'  ^^  ,i»,-,,i 


iSKsaw^  -i*»*. 


Post  root 
Ant  root 


Preganglionic  fiber 


\      Sympathetic  ganglion 


Wfijfe  rami 

Postganglionic  fiber 


>'.     .I'M;    »l.i.  h    .; 
'    .--     ■:    T      f.rm     11^ 


if  .¥i 


k,-  -,    :  I 


.  I 


11    i 


Till:    M    inMlMII      SI  UMll  S    SVMI  \l 


SHI 


sl.llatiiiii  for  111''  fiiif  liiiili,  ami  tiir  lnwcr  limibar  ami  iippi  r  sai-ial 
'.MM'.'lia  I'm-  tlic  liiml  liiiili.  Wlicii  tliry  air  ^linnilatfcl.  tin'  niiisciilar 
tillers  MiiiiiiiiiilihtJ  tlio  sweat  u'lamls  conlrnct  ami  srniecze  out  tin   sweat. 


'-Ill    MMiN  UK  lll.DOll  Sirr\TMiN    Of    MOTOR 

vrssKi.s  (, AM, I. ION  et.i.i.s 

Mr:!. I  :iii'l   Mcc-k.  Sii|ii  lior  I'crviciil  i;:iii;,'Ii"n. 


Il.mt. 

Alifilinl    lAtlflllitv.         (;;ili;;liull    «lrll;itliril. 


(laiiulinn    xli'lliitiiin    iiikI    in 
fc'iiiir  ciMvii'til  can^Iiiiii. 


l;i>o|  -i  1  n\  1  MMM,    (  clNSI  (    IliU 
Nl  liVI  S 
1.  '2.  ;;.    I.  'i,  ll-  tMri.-;  L',  .'!,  ■»,  i;ivi' 
iii;i\imiMii   ('IT«M-t, 

1,  :;,  :',.  i.  r,.  ili..i;,,i,-  2,  ?,.  ^h.- 

niaxinniiH. 

I.  .'i.   li.    7.    ^,   ;i,    U.'n:ii-ii'.   !inil    ]" 
>li:;litlv. 

C.-trriMi-  cxticniily.      (Iili   li.inl.Mi-,  Till  liiMil.:ir.  iiifl         H.    1-.    1  ;.   ll>"iM,i,- :    1.   L'.   lninl.:ir 

lic.iMl  i.';ni«li"n.  I.    .",    <:.    T.    V    '■'.    1".    H.    1-.    l'. 

tliiiiiM'i.- ;    I.   J.   .".,    I.   liiiul>:ii-. 


Ki.liicv. 

>|'l(rll 


Si-inilMii:ir  ;::iM^liiiii. 


.     t.    ."..    Il,    7.    >•.    '.I.     Id.    11,    1L',    1'.. 
ilni.i.h- :    I.J,   '■'..  Iiiiiiliiir, 


.\I-I,,iiiin:i|   \isci'r;i,        Sii|ur|iii   jiicMntnic  ;;iiii;;1iiiii        <'■    7,  ■<,  l'.  in.  11.   i::,  llini; 
.■111.1   si-iiiiliinnr  ;;iin(ili..n.  :'.   Imiil'Mr, 


I'l  l\  il'  \is('('i:i. 


I.  !iinrl.;ii. 


.  (i^.^K.in 


The  <^aiii;lia   fur  the  iiihunolur  jUk  r.^  are  ninre  \\  kli  spreail  iestoiuliir-' 

I'luiii  the   I'niirth   thoraeie  to  the  evL'oal   jranu'lin"! ;  hut  the  eonnoeti'r 

liheis  arc  au'aiii  strii-tly  e((iitiiie.l  to  the  llinraeieohmihar  logi'ni,  Sfiiiiu- 
lalioH  of  these  fillers  eauses  iiiovemeiit  of  the  hairs,  or  on  liairloss  aiiiiiiaN, 
the  (•(iiulitioii  called  "  y;oose  skin." 

The  Motor  .\iiris  ot    Ihr   Miischs  Siirroiniiliii'i    iln    Si  t/nn  iiln!    Ihirl. 
It   will   he  ohsiMvcil   that    the  ci>ntiri-tor  lilicrs  to  the  alHloiniiia!    aid   pel 

\  ie  \  isccra  an llected   into  two  special    iiei\-e  trunks,  the  L're.-itcr  ami 

tlio   lesser   (or  lumhari    splanehiiics.      The   collateral    •.'alidia    (semilunar 

aiicl  snp<Mior  ami   inferior  iiieseiitcric^'    wilh   which   these niiect,   lia\c 

iiotliiiii.'  to  ilo  w  ith  the  Niiiiiiciital  mMM's,  Imt  their  ner\e  cells  sciid  (iliei's 
'  postiranjilioiiic''  which  siippl\  the  \arious  viscera  not  only  \\itli  \as.- 
iiiotor  filii^rs  luit  also  \xitli  the  '•  s>  lupatln.'tic"  (ilieis,  which  w  i'  h.'Ui'  seen 

exercise  siicli   an    imporlaiil    itrol    o\cr   llicir   ■_'i,iiiilii!ar   ami    iiiusciihii 

functions. 

All  of  the  fillers  .•uiilaineil  in  the  liiiiili;ir  splanehiiics  do  tmt.  ImwcMT. 
h.ne  their  cell  stations  in  the  inferior  iiiesentcric  liaiiu'lia.  I'lil  riii! 
thriiil^rh    it   ,iiid    proceed   in    the   li\  poi;astric   nerves  to   tind    llnir   elVietdr 

eells    nil     llie     111  llseuhlt  11 1'c     of    the     various    si  lllct  1|  l-'s     th.lt     a  r-     .I.Mlnpi-ij 

fruMi   Ihc   .Miilleiian  and   W'olffi.-in  iliicts      i    c.  of  Ihe   ureters,   iit.riis.   p'al 


i  ;i 


•  \i 


8H2 


Tilt    CINTKAI.    StKVOI  S   SYSTKM 


lopinii  tiilips  iiihI  vns  (l.'lVicns.  Mjuiy  of  thf  HIkmh  i.l"  the  livponastrif 
iHTvrs  iiiT  tliPiTt'oiT  those  of  involuntary  iiitrrniniciiil  lUMirons. 

Till-  Ihiiiiilir  iOiil  iiiliniiil  mini  sphinrti  r  i.ii/sr/f.v  of  tlic  intestines  iind 
iiitirml  visiiiil  siihinrlir  reeeive  their  nerve  supply  from  effei-tor  neurons 
situated  in  the  superior  and  inferior  niesenterie  na>>!.'l'i'.  ''>*"  intei  iinneial 
tihers  arisini:  from  the  thor.ieii'olundiar  rei:ion.  It  is  |)ossihle  that  the 
other  sjihineters  of  the  intestinal  eanal  viz,  the  eardiac  and  pylone 
spliini'ters  of  the  stomneh     are  similarly  iniu'rvated.     ( l-'ij;.  '2:i2  i 

(ireat  aid  in  workinir  out  these  nerve  eonneetions  is  reeeived  liy  stud\- 
insr  the  etTei-t  of  i  ;  iiiephrine,  wiiiidi  arts  speeirieally  on  those  tiss  u-s  that 
are  supplied  liy  I'le  symi)athelie  nervous  system.*  Kpinephrine  has  no 
elTeet  on  tissues  iiuu'rvat.'d  hy  the  hulliar  or  saeral  outtlows.  and  it 
develojis  its  aelifui  ))eripherally,  lieiiij;  imleed  more  potent  on  a  dener- 

vated  orvran  even  after  all   its  nerves  have  1 n   alio-,  ed   to  deu'enerate. 

Advanta^ie  of  this  aelioii  of  epineplirine  has  lieen  taken  in  the  itnesti- 
•jalion  of  doulitful  eases  of  sympathetie  iinu'rvation,  sueh  as  in  the  eere- 
Itral.  eoronary.  and  pulmonary  Mood  vessels.  The  outeouu'  of  these  in- 
vestiiiations  has  lieen   diseussed   elsewliere. 


■1! 


[%r 


THE  BULBOSACRAL  OUTPLOW  OR  THE  PARASYMPATHETIC 

SYSTEM 

I-'roiii  the  MU'dulla  olilonuata  arise  involuntary  eonneetiu'  neurons, 
uhieh  are  earried  mainly  liy  the  vairus  nerves  tuit  partly  liy  the  seventh, 
ninth  and  eleventh  eranial  nerves  to  elTeetiu'  nerve  eclls  situated  periph- 
erally on  the  struetnr-es  to  whieh  the  nerves  run  (  \'\'l.  'I'-V-iu  These  include 
in  a  ueiieral  way  the  museles  and  irlands  of  ihe  alimentary  eanal  an.l 
its  derivatives  as  far  as  the  end  of  the  snndl  intestine,  in  the  simdl 
intestiiu'  itself  the  cells  of  these  motiir  tn^urous  are  those  of  AiU'rliach's 
]ilexus  found  lietween  the  tuo  muscular  coats.  In  the  diverticulii,  which 
include  the  lunsrs  and  the  trail  Madder,  the  nerve  i-ells  to  which  the 
\  ajrus  fillers  run  are  also  situated  peripherally. 

The  smntl  ititlpoiv  occurs  tlimu^rh  the  seeoml  and  third  sacral  nerves, 
the  liliers  .joininu  to  form  a  single  nerve  Mhe  pelvic  nerve  or  nervils 
eritfciis;  on  ench  si.lc  This  i)asses  directly  to  the  Madiler,  where  it 
connects  with  a  plexus,  often  called  the  hypotfastrie,  which  extends  over 
the  Madili'r  ••mil  neiiihliorinjt  portion  of  the  rectum.     The  tiranchcs  run 

1, nnect  either  with  the  nerve  cells  of  the  trauiilia  of  the  plexus  its-.'lf, 

or  with  nerve  cells  situated  on  the  walls  of  the  larijp  intestine  and  Mai'.- 
der.     The  jielvic  nerve  iiudies  its  connections  with  the  periphery  in  the 

•h«  arti'^'i   i>  al*;r.«  tlir  «.T:ic  .1^  Ih.if  wHuh  is  i.r...|u.  ril   l.v   s'iM,uhili..ii   "<    tlic-   »\niiallM  iw     miw 
■  upi'ly.  whcthiT  tlii>,  rffi-.t  IV  i.nr  ..I   «iiiiiulati"!!  "t   mliilHIi'm 


■*^„F'*"-^/i"i.-.i«' 


^    —   Key  - 

reCan^iionir  Syinp<jtlietic 

■  e  (ianglionrc  Bulbo  Sacrjl 

Pt'ii  uj'  ji'  >■  'v  n  I'r  !  i.!  '  ,11 


Heado 

Neck 

Thofjcic 

Coelidc  Pieiuifr 
Jup  Me]  Cdn^ 

Leg 


Arm 


Abdominal 
Vijcer« 

lPr»()«n()J 


Lumlur 
Spidiiciiiiic  nerve 


"i^  _i  —  Inf  Mej  Gang 
i 


/i.if 'A 


ASp.  irt. 
Peine  visceral  nerve 


«'rti      t  >^        *:'  ti         tilt      'ik.      ..I     i'.ilir\      ^'Vf.i     ..1     tt  . 

i-ll'Mi--i  h  -    .iri-     .i:-:itt«il.     f.-.it     llu      ].ii»it!Mi-     ..t     tii.r     ,Bi.^^     Ii-'llt     'it 

'I..I.-  111.     .1-ui.ll"     .l,...v<.    .  l..,lK     111.     ,iF-liil-Ml.-,li     ..1     II 11...- 

i.|    Ih.     V.1UM-  .li.-l    ih.     i.l't.   -,.■■!,.!    .111.1    !lnl.l    -..-  ■..;    i.iv- 


\..-.l^     ^^-ti-;..     t.t     ).»      11.11    I     .it.. lie 

,         1.1. .       -       .1       111.         .1111.. .III.,  lu 

.  ..f.l    1-    in.ti.  Ill  .1    II.    ill.     .ii'lt 


^rl 


M   :! 


Tin:    .\l  TtlNOMIC    NKKVOI  H   KYKTKM 


hm:i 


snnir  ninniM'i-  as  tlic  villus.  Tnkcn  foKcthci'  tlicw  t\Mi  m-rvt's  nupply  tin* 
iiiUMc\ilntui't'  of  tli(>  KaHtriiiiitcNtiiial  tract,  iiicliuliii|i  tin-  cloai-a.  the 
vamiH  ns  far  an  tin-  cnil  of  tin-  small  intt-stino,  and  the  |n'lvic  hir\f 
fforii  tliis  point  oti.  It  rmist  "f  coiiisc  In-  ri'inonilx'rTil  tliat  I'l-i'tain 
imisclt'H  namely,  tin-  >pliin<-tt'r.s  of  I  lie  Kmall  and  laiKi-  inti'stinc 
riT.-ivi'  tlii'ii'  n<T\c  Kiipply  fmni  tlii'  MvnipatlH'tif  (pairi'  XX'2).  -Inst  as 
<<li  ui'liiri's  iiiii(f\  ati'il  Ity  the  «>  Mipatlictic  art-  pt'i-iiliarly  Hiix'fptil'l''  t" 
tlic  action  of  ('pini>plirini',  it  lias  lircn  i|isi'o\ci'i'>|  tjiiit  tliosi-  inncivaicil 
liy  tlic  liiilliosacral  system  arc  very  siisi-cptililc  io  tin-  action  of  acctyl- 
cliolinc,  which  is  prcsriit  in  cit'ol.  Tiny  arc  not  acted  on  li>  cpinci)liiine, 
nor  are  the  siructnies  npon  which   ilus  acts  alTected   liy  acct\  j-clioline. 


AXON  REFLEXES 

At  this  pl.iie  it  is  eiin\rnient  to  consider  for  a  nionieiit  the  phenome- 
imn  which  has  alrciidy  licen  referred  to  as  an  axon  rcllex.  It  wa'*  dis- 
covered tliat  when  one  of  the  hypoi^astric  nerxes  was  nit  and  the  central 
end  stiihulatcil  there  was  a  rellex  contractoni  ot'  the  liladilcr  anil  the  in 
ternal  anal  spliiin-ti-r',  alonir  with  vas iistriction  in  the  r<'tiion  of  the  rec- 
tum an.!  that  tin-  cic.ui'ieil,  even  after  diseonncetiiii^  the  inlVrior  mesen 
Icrie  uMiiirlion  fnun  th,  spinal  eoid  liy  cnltiiii;  lln'  hnnliar  splani'hnii'  nerves. 
Injectii'i  of  nicotine  iaimediately  idiolishc(|  the  response.  It  looked  as 
if  rellex  action  was  |>risNili|e  tlironirli  the  lianirlion;  which  wouM  jnstif\ 
the  name  "syinpalliet  ii"  oriirinally  ifiveii  to  the  involuntary  nervous  sys 
tern  in  the  helief  iliat  the  iranitlia  were  ei>ntcrs  foe  loeal  rclh-x  actions. 
Further  iii\  cvtiLration  slmweil,  howcxcr,  that  this  lellex  is  not  similar  to 
those  oceurririL'  in  the  \oluntar>-  system,  Imt  is  dependent  npon  the 
presence  of  a  <'olliitcral  on  intcrnuneial  (iliers  that  run  tlirouLrh  the  in- 
ferior im'seiiterie  (riitiirlia  to  ner\c  cells  situated  periplierally  on  the 
Malls  of  the  liladder  and  rectum.  The  collaterals  terininate  hy  synapsis 
jirouiid  nerve  cells  in  the  >rany;li<'n,  the  axons  of  which,  as  we  have  seen, 
rini   to  the  liladder.  the  rectal   lilooil    vessels,  anil  the   internal   sphincter 

ani.     Tl videnee  for  this  explanjition  depends  on  the  oliservation  that 

the  axon  redex  is  no  lonjier  |)ossililc  iifter  the  linnliai-  splnnehnics  have 
lieen  cut  and  time  allowed  for  their  tiliers  to  liecome  eompletely  deuen- 
erated. 

Similar  rcllexes  ilc|>endinK  on  eollateiais  have  lieen  found  in  the  lateral 
chain,  and  there  can  lie  little  doulit  that  they  arc  of  fre<(uent  e,<-eurrence 
throutt'iout    the    whole    involuntary    nervous    system,    just    ;is    they    are. 

within  the  spinal  cord,  in  the  voluntary.     It  is  hecause  of  thest llatcrals 

ami  the  fact  that  in'rve  liliers  transmit  impulses  in  Imth  <lir<'ctions 
that  a  stwiiidns  transmitted  throiinh  one  or  a  limited  numlier  of  connect<ir 


MICROCOPY    RESOIUTION    TEST   CHART 

ANSI  ond  ISO  lESI  CHART  No    2 


1.0 


1.25 


|-I2  lis 

t  !f  IS 

1.8 


14  mil  1.6 


A     APPLIED  IfVMGE     Inc 


>sl 


iiii:  riATUM.  Ni  uvors  sv>ri:M 


iiciiriiiis  iii.iN    cM-itt'  a  biii.iil  lii'lil   of  ('(Tt'i'lors  aiul  cause  a   \\  iilcspread 

.  iVirl, 


lU 


'  { 


JNCTIONS  OF  AUTONOMIC  NERVES 

'I'lie  fiiiic'tiiiiis  of  tlic  aiitdiKiiiiic  ii('r\e  lilicis  liavo  Ix'oii  discussed  in 
(•(PiiiiiM'tiiiii  \\\\h  tlic  st  iiictuit's  wliidi  llu'v  sui>!>ly,  and  we  sliall  ro(|uiri'  in 
ill's  iilace  only  to  review  llieiii  in  a  trenei'al  way. 

Two  opposed  ctTeels  niav  lie  nlitaiiiiMl:  stiwnliildrii  i  auiriiieiitoi'v)  and 
!iiliihlfor>i :  and  llieso  may  lie  ]irodiieed  tliroutrli  one  nervp  li.\'  it;-'  l)ein<ir 
stiiMtdatory  for  one  set  of  niusele  fibers  and  inliibilorv  foi-  anotlioi'  set 
ill  tlio  same  \  iseiis.  Tlio  liianelies  runnin<;  from  tlio  inferior  mesenteric 
uantilion  to  tlie  colon,  for  example,  are  auymentory  (constrictor)  for  the 
lilood  vess(>ls  and  iiiliiliiloiy  for  tlie  muscular  walls  of  tlie  colon. 

The  frrcatest  interest  centers  on  the  inhibitory  impulses.  They  arc 
lust  known  in  einmcetion  with  the  vatrus  nerve  to  the  Iwart,  the  synipa- 
ilietic  to  the  >m;ill  iiilestine,  and  the  hypufiastric  to  the  miisculattire  of 
tlie  bladiler.  It  is  iiilei  (stiiii;-  to  cnmpare  the  nature  of  inhibition  in  the 
iii\  (limitary  ,ind  Milinitary  systems.  In  the  lattei'.  it  will  bo  rcnienilicrei'l, 
inliiliitioii  (■,•111  iicciir  mily  tlirouuh  the  intcnmneial  neurons  and  the  ef- 
I'eilipr  nerve  cell,  stimulation  of  the  efl'ector  nerve  fiber  never  haviiij: 
any  other  than  an  anirmeiitor  elTect.  It  is  tpiite  otherwise  in  the  involun- 
t.iiy  iierxniis  sxsteni.  fur  stiniiilal  ion  of  the  elTector  nerve  fd)er,  after 
niiiiplete  destruction  of  the  cfTectur  nerve  cell,  is  still  followed  by  a  typical 
iiiliiliition.  This,  jt  will  be  remembered,  nniy  be  denioustrated  on  the  frojr 
hi  art  by  ap|il\  iiiir  elect  lie  sliiiiulalion  to  the  white  ci'cscentie  line  after 
pariily/in<;  tlu'  effector  ner\e  cells  liy  nicotine.  The  same  may  also  be 
^liiiwn  in  the  case  of  the  chorda  tympani,  wlii're  stimulation  of  the  )iost- 
'.^.iimliiinie  fibers  in  i1h>  hiliis  of  the  irland  causes  dilatation  of  the  blooil 
\isscls  after  jiaralysis  of  the  fxanfrlion  by  nicotine,  vasodilatation  beinir 
iif  ciiurso  a  phiii-'uiennu  of  inhibition. 

It  is  a  difticnlt  matter  to  desiirnate  precisely  which  fibers  in  any  i)ai't 
nf  the  in\  iiliiiitar\  iierMms  system  are  inliibitory  and  which  auirmontory. 
Indeed,  as  mentioned  abo\e,  one  liber  may  pei'form  both  fuin-fions.  In 
eases  where  the  exisfeiice  of  inhibitory  (ibers  is  diHibtful,  iirrcat  aid  is 
affiirdi'd  b\  the  Use  of  ei'irotuxiui'.  an  alkabiid  uf  ecLiot.  wliii-h  |)ossesses 
the  I'emarhable  property  of  specifically  paraly/inir  the  auuniu'iitor  nerves 
nf  the  s.\  mpatlK'tic  system  'but  nut  of  tlie  iiai  asympathetic'* ;  that  is, 
ilir  s.ime  fibers  as  are  stiiiiiilated  by  epinephrine.  When  a  particular 
st  riicl  me  is  snpplieil  \\itli  .'imrtiientory  and  inliibilnry  liber:,  by  ii  com- 
iiiiiid  >.\  uiji.iiiii  '  ie  i;ir\e,  ,-iictrie  st  ininlation  •■]■  tin  .•ipp!ic;if  ion  of  epi 
iicphiiiie  Usually   t.'i\rs  oiily  an<rmentiiry   elTecIs;  after  the   injection  of 


l^i ' 

1  . 
1  i 

-mi  '"'  ■         '    ;     i 

SI '  i' 

■< 

Si ''  I 

'        ■'■ 

^Bl 

^il 

ikk^ 

\pilo  motor  i.iu5C/e 


Lachrymal  q^and 


CilHryging      ^^ 

Uasalmucoia  ^  \_) 

SphenopaliTaa 
Sublingual  /\        ~    Parotid 
gland    [    | 

'"-^-V — ^  ^'V^ 
Submaxillary  glanC  \         \/ 

6ubmaxillarY(5ublinguai)    ^^ 
ganglion 


Lung 
Call  bladder  and 
ducts^,^^ ..__^ 


I    Cranial  and  Sacral  nerves 

'^  I  inhititory     blue 

^r*^       [Ttioraclco-lumbar  nerves 

Postganglionic  fibers 
are  dotted.thus-  - 


N.XI 

Sup.  cervical  ganglion 


3mall  infest. 
Pancreas 


lliocecal 
sptiinr     r 

B/a3der 

Vesical  sptiincter  J 

Urettiral  sphincter//, 


Thyroid  gland 


Inf.cervical  ganglion 
T\j;-Ansa  subclavia 

Stellate  (I'J  Thoracic) 
<A  ganglion 

'^^t  Sweat  gland 


^  Vasomotor 
fibers 

Pilo  motor  muscle 

—  Celiac  (Semilunar) 
ganglion(Solarplex) 
Splanchnic  nerves 

Sympathetic  ctiain 


[Lumbar  splanchnics 

Sup.  Mesenteric 

ganglion 


—  Inf.  Mesenteric 
ganglion 
Hypogastric  nerves 


Pelvic{HYpoqa5tric,interiliac)   (  sphJncier  , 
plexus  (Vesical arectal portions)      Pflvic  nerves  {Nervus  erigens) 


P    r  HdilvcK 


" 

1 

1" 

1! 

w 

;      l) 

:j!i: 

- 

1-ii 


it ; 


'U 


I  > 


I 


i'l  il  i 


.!  :       t 

•1 


i! 
n 

^y 

] 

if 

1  ^ 
* 

1 

1 

n 

r' 

Tin:    AI'IdMiMIr    MHVdl 


>vsti:m 


8sr. 


(•r(j;o|(ixiiio,  Ikiw  cv  ir,  ji  icvcrsfd  cITrct  is  (ilis(  !\  cil ;  iuiiiit'l>',  iiiliiliiliim 
iiistond  of  iiiifiiii<'iil:i1ii(ii.  M.v  liikiiiir  iiilviiiit;i'_'c  of  lliis  I'.Mi't,  Hale  lias 
licrn  iilili-  t<(  (Icnionslriitc  in  llii-  liypouasliic  nerves  inliihitmy  (ihors  to 
tlie  iileiils,  and  Klliolt  lias  (leni'instiateil  tlic  inliiliitoiy  aetiun  ol"  epi- 
iM'|(lirino  on  the  muscles  of  the  iiretei-  in  llie  doir.  Inliiliitoiy  lihoi's  liaM' 
also  lieen  diseovered  liy  lliese  inetlio<ls  in  the  LM-eat  s|ilaneliiiii'  nerves,  in 
tlie  neive  roots  siiniilviiif;  tlio  kidney,  ;ind  in  the  eer\ii'al  synipallietic 
s\ipi)l.\  in;r  the  lihuxi  vessels  of  the  nmeoiis  nieiahrane  of  the  inoiitli,  ete.; 
that  is,  in  nei-vo  trunks  which  i)revious]y  were  lielieviil  to  <'ontain  only 
ansinicntoi'y  filicis.  The  acconipanyinLr  diaj;'ain  from  .(.lelison  will  pixe 
an  idea  of  the  cni-rently  accepted  xiews  concernini:  the  distriliution  of 
aiiynientins    and  inhilijtory  lihers.     (  l''i'_'.  2'-V''t.> 


THE  AFFERENT  FIBERS  OF  THE  AUTONOMIC  SYSTEM 

It  has  loiiy;  heeii  known  that  the  exposed  viscera  are  leniarkahly  insen- 
sitive. This  experience  is  in  acci  d  with  the  oliservation  that  the  siii)i)ly 
of  afTcrent  filters  to  the  viscera  is  relatively  very  small.  In  the  hypo- 
gastric and  probably  in  the  irrcat  splanchnic  nei'xc.  I.anu'lcy  computes 
that  oidy  about  one-tentli  of  the  medulliited  fibers  are  atVei-enl.  At  tin- 
two  end.s  of  the  aliment'iry  canal,  where  cooi)erati\ c  rellexes  between 
the  somatic  museidature  and  the  viscera  are  of  inipoitance,  a  ".neater 
numbei"  of  afTcrent  fibers  are  foimd  in  the  autonomic  nerves;  fur  ex- 
aniide,  in  the  pelvic  nerve  about  one-third  of  the  fibers  are  afTcrent,  and. 
as  we  have  freriuently  seen,  the  vagi  contain  large  nnmb(>is  of  tlieni 
coming  from  the  lungs,  stomach,  and  no  doubt  from  other  viscera. 

Tlie  afferent  visceral  fibers,  as  abov(>  stated,  arise  like  those  of  tin' 
voluntary  system,  from  tlic  ganglion  cells  of  the  jiosterior  roots.  The.\ 
travel  in  company  with  the  cdnneetor  fibers  through  the  white  raiiuis 
communicaiis,  so  that  the  stimulation  of  the  central  end  of  one  of  these 
may  cause  I'cflex  rise  in  blood  i>ressure  and  oilier  movements. 

It  is  found  that,  after  o|)(iiing  the  abilnininal  cavity  undei'  local 
anesthesia,  cuttinu'  and  sutui'inir  of  the  \  iseera  may  be  enntinued  without 
causing  any  i)ain.  \Vhen  the  \isceia  ari'  inflasiied,  howevei',  and  umlei' 
ccitain  conditions  of  stimulation,  such  as  the  distention  of  the  bile  duel> 
with  biliary  eabiili,  or  the  violent  con1raeti"n  of  the  intestines,  excruei- 
atiiiir  ]>ain  may  bo  evokeil.  This  pain  is  fre(|Uently  not  localized  in  the 
\  iseera.  but  is  referred  to  certain  i>ar1s  of  the  surface  of  t!ie  body,  and 
it  has  been  shown  by  jra(d<en/ie  and  b.\-  Head  that  it  is  i-eferred  to  the 
area  of  skin  which  is  sui>plied  with  sensory  nervi's  by  the  same  segment 


tliiit   to  \\hieh  the  nfTereiit   autonomic   (ilicis  o 


>f  Ih 


p:i! 


run.     It  has  further  been  shown  that  xasculai'  disease  ma\    cause  sens! 


i  1.  i  i 

li 

V-:\ 

SSfi 


Tin:    CKNTRAI-    NKRVOfS   SYSTEM 


tivity  tit'  fill'  ('(iricspoiuliiin;  cutaiicous  ari'us,  so  tliat  cliiiieul  methods  arc 
available  for  localiziiijr  tlio  site  of  the  disease  l)y  studying  the  exact 
jHisitioii  and  extent  of  the  I'eferred  pain  or  skin  tenderness. 

XHRVOrs  SYSTK.M  RKl-'KHKNTKS 

(Monographs  and  (iri(;iiial  I'a])ers) 

'I'iirkiT.  <i.   II.:      \'\,n-.  Am.  VhWos.  ^     .,  lull,  i,  ::i7l':^.">. 
-il.'ii.l,  I!.,  iiii.l  WwnH.  \V.  II.  |{  :      l^i^iii,  l',i(is,  x.xxi.  .'ilili^.")!!. 
Mirk,  \V,  ,1.:     Am.  .Icmi.  I'li.\siii|.,   I'.iH,  xx\iii,  ."..Vi  .".ilo. 
iliiuii',  A.  Xiniiiii:     Arcli.  f    rxi^cr.  I'alli.  ii.  I'lmrmiikol.,  llUu,  |.\iii,  4l'tl  t:i.!. 
'■'SliiTiiimtiMi,    <".    S. :      Nuiiiiiiiii.-    |>ii|ii'is    III!    iciipripi-iil    iiinrivatiipii    ol'    iuitii;;i>ni»tic 
iiMi.srh'^t,   I'lur.  Kiiv.  .Sell'.,  \i)l.   H,  (ili;  ulsii  ill  .liiur.   I'li.vsiol.,  xxii,  xxxiv,  xx-viii, 
xliii.  ami  (jiiart.  .Iiiiir.  KxpiM.  I'livsiol.,  ii. 
■lliilnics,  (ioiiloii:     Hrit.  Mid.  .lonr..   I'lM,).  ii.  Xi.v.  1'7,  !).•.■.    1  an.l   11. 
'I'iki',  y.  11.:     .\m.  .Imii.  I'ii,\>i(il.,  Umiii,  xxiv,  iL'll.'iJ. 
Miillv,  \V.  .v.:     yiiait.  .lour.  Kxiut.  I'livsidl.,  I'.tlll,  iv.  (17  ^7. 
M.Mml.ai.l,  W.   1'.:     .lour.   IMnsidl.,   iss'ii,  x,   1:!^  Us. 
'•(•..Ili.T,  .1.:      I.amrt.  April    1.'  l!U(i,  711. 

"■Kaiis.111,  S.   \V.,  iiiiil  villi    Hess,  C.  L.:      .\iii.  .Imir.   I'li.vsinl.,  liil.'i,  xxxviii,  12s, 
"Ilcail,  II.,  aiiil  Thomiisiin:      Ilrain,  liKlti,  xxix,  ri:;7. 
i-Sln  rriiiHtiiii.  C  S.,  ami  Hniwu.  T.  (ira)iaiii:      Jniir.    IMivsiul.,   I'.'l.'!,  xlvi,   Pnir.   I'li.v.siiil. 

Sill'.,  |i.  xxii. 
1   Itnmii.  T.  ('.Taliiirii,  ami  S!ii'i-iiii;£tiin.  I".  S.:      Pine.   Rnv.  Siii.,  WtVl.  s.'i,   H,  J.'ill  L'77. 
"I'lisliiiiy,  Uarvi'v:     I'ror.  .\m.  I'livNinl.  Siir.,  .\iii.  .tmir.  Phy.siol.,  ISiOli. 
i''l.iniaiii,  I..:      Ivli'inliirii,   Krurliiiissr  iIit  Pliysiiil.,  1!I(M,  i. 
1' lilark.  Daviilsiiii:     <'i'r('l«'ll!ir  I.iirali/alinii  in  tlii'  I.iulit  iil'  Koi'cnt  Roscarcli,  .Iniir.  I.ali. 

ami  t'lin.   Mi-il.,  I'llii,  i.  4117. 
'■(laMki'll,  \V.   II.:     Til.'  Invuliiiitarv  Neiviiiis  System,  Miiiiiiaia|ilis  on  PliVNiolouy,  eil.  Iiy 
I-;.  II.  Sttiiliiii;,  l.iinyinaiis.  (ireen  &  Co.,   l<IHi. 

Other  J[i)iKii;raphs  not  Specifienlly  Referred  to  in  the  Context 

''Sheninntiiii.  ('.  S. :     i  I  i   The  Ititi'^iative  .Vi'timi  iif  flie  Nervous  System,  Siliiiiiaii  Lee- 

tiiris.   Vail'  riiivirsity.     Srrilnier's  Sons,  New  York,      i'l)  Sliiifer'.s  Te.xtii'nik  of 

Pliysiiiliicy.  II.     Yimi'iL'  ■!.  I'entlaml,  I Inn,  ISilH. 

'"Hiiltiiii,  .1.   S.:      Ki'ieiit    Keseanlii's  in   Cortiial    Liii'aliziition   anil   on   Tlio   K'lnilion   of 

tlie  CinOirmii  i:.  Fiirthi  r  .liliiniri a  if   I'liiisiiititjiii,  eil.  liy  [..eonaril   lliil,  F.m.iion, 

K.  ArmiM,  I'JO'.t. 


INDEX 


AlKliiniinal    ri's|>iratiiiii,   'M>7 
Ahiicinnal    imlsis,   JTii 
Alisiirptioii,    in    ({'''"''""'i    '•' 

from  Mtoinai-li,  4~i>) 

<if   fats,  •)!•! 
Acapnia,  :!!•-' 

Acrt'ssiiry   fcidii    facfnrs,  ."84 
A"cfal(li'hyclc,  T"H 
Aiftciacotic    aii.l,    fiS.!,    7(l!l 

Airt ',    tiS:(,    7(i!t 

A.i.l: 

liufTiT   aiticiii,   :fti 

I'xrri'tidii  (if,  liy  kidneys,  4t! 

niiMilicr    iif    fats,    tiH7 

tcital   conci'iitration   of,   .'i:! 
Acidity,  actual   dc'tjrcc   of,   '2'.\ 
Acitlosis: 

ammonia  urea   ratio   ilnrin^;,  (llii 

compcnsatod,   .'!!• 

in     diabctfs,    tiH.! 

in   ni'pliritis,  tiH:t 

in  starvation,  ."i(i9 

rolationsliip  to  alvccdur  ('(V,  .'i.">4 

ridationsliip  to  linatldn^,  ;!.~i4 

theory  of,  :ts 

nncompi'iisutcd,    .!!» 
Acids,  of   nrint',   ')'24 
Actual  df^ri'i'  of  acidity  and  alkalinity, 
Adenine,  ti;irt 
Ailenosine,   *»;»H 
Adjusters,    "s:; 

Adrenal  glands  and  dialiites,  (i7;t 
Ailrenalini'    (.st<     Kpincplirinei 
Adsorption,  li'i 

ciimpoumls,    70 

ciuKlitions    inlluence<l    liy,    (i7 

effect  (if  chemical  forces  on,  (>H 

effect   of  (dectric   tdnin^jes  on,  G7 

everyday  reactiiuis  lUpendiiiK  (Ui,  tit) 

of   biases,  *>*> 
AflFerent  fibers  of  autonomic  systi'ni,  HS.'i 
Afferent   spiinil   pathwavs,  HIill 
AKe,    r,,S4 

effect    tui   creatinine   excretion,   fi«f 
Alanine,  find,  (W.i,  tiuCi,  til!),  tititi 
AUiolene   absorption,    til>l.' 
Albuminuria,   fiH) 

Alkali  retention,  determination  of,  48 
Alkaline   buffer.  .'Iti 
Alkaline  reserve,  'M 

measurement   of,   41 


Allantoin,  li.'tli,  li;i!t,  t'>4.'i 
Allied     reflexes,     simultaneinis     intejjration 
of.   hlM 
successive   integration  of,  H-ll 
Alloxan,  ti.t."> 
Alve(dar  air: 

clinical   investi^atiim   of,  .'!47 
estinmtion  of  ^ases  in,  ^'44 
Kridericia  method,  .". ID 
Haldane   metlioil,  ;i4<P 
I'earic  method,  :!4.") 
tension  of  ('()_„  4ti,  .i.i!*,  .ir.ti 

durinj;    breathing    in      ontineil    space, 
.•!o7 
tensi(ui  tif  oxyj^en,  X\[* 
Andiard's  e<|nati(m,   .■i;;7 
in  acid  excretion,  48 
And»oce])tor,    '.Mi 
Amino  acids,  .'ii»7 

and  eniT>;y  output,   ')4l 
in   blood,  tidti 
chemistry  of,  '}'.*H 
iletermination  of, 
fate  of,  (ill) 
Krunps,   ."lilH 
in  (jniwth,  .">7t> 
in   tissues,  ti07 
in  urine,  ."ilUl,  (iL'ii 
struitnre  of,   lillL', 
Andnoacetic  aciil    (,s<c 
Anunopropionic  acid    (mc  Alanine) 
Annnonia: 

amnniuia  urea   ratio: 

inllnence   of  acidosis   cm,  (il6 
in    clisease,    t!:.'il 
inffiu'uci'  of  liver  on,  017 
as  rcservi'  alkali,  tilti 
excretion  of,  til.") 

excretion  cd'  ;icid  in  combination  with,  46 
of  urine,  .">.'lo 
.\niuinnium  larbanutte,  fill! 
.Vmmonium    carbonate,   tiltt 
.\nioeba,    78:.' 
Amylases,  ,H1,  (Id,  4!II 
.\mylolysis,  4111 

in  stonuuh,  4.14 
.\mylupsin,    4!ll,   6.">6 
.\riacrotic  wave,  pulse,  !20:i 
.\nalysis    (psychic),    S.")8 
.\naphvlactic  reaction,  ."I'.i"),  001 
\iirip!iylaxi»,  8H 
.\narthria,    8(iL' 
Anastomosis,  intestinal,  470 


"liM.t 


tiO;{ 
lilycocoU ) 


i                ■                      ' 

1 

W' 

%\: 

\1,.M1L1.     '.'.'. 

l.l,H,.ltl"»    ill,   ■-•><:: 

\lit>llir-i:i.    h:;! 

Aihiiri«iii,  1.1 IiImiv    ill,  L'sl 

IuiIm'  ill,  1 1::,  :.'  "I 
\ii«iii:i    i...t..ii«.   Iil.rill;,li..ii    in,    l!"l 
.\iiitpi;il    cali'i  iiiu'tir,    ."i'.ii 

\Mii.iis,  ir.,  :.:i 

\iit:i^i'iii^ti>-   tiiii^i  Irs,   sis 
Aht:i;;"iii-li.'    hiIia.-.,  s.' I 
\nlrri.ir  niuls,   7s7 
AliIiio;iKlll:iiits,    !'!• 
\nti4riiiiiii-    iiiiiMiNi's,   lMI 
AiililViiiiiMiti  ill   I1I1H..I,  Ml 
AiititliTonil.iii,    Iiil,    II- 
\htil"\iiis,   ii',1 
Aiitilrvpsiii,  IMI 

Anrti.-    iijjiiijjitalion,    piiUr    in,    \-''\ 
A|M  ~tlic<iii.  s:;h,  s:,i 
A|..\   l..:il.  Iniiiiiu  i.r.  -ll'i 
A|lia~i:i,    iiH.tnr,    siln,    si;j 
M'tisitiy,   Miu' 
Mili.iiiii.-;il,   x>'<- 
A|ihi':l,   llrlViilK  llillirlll    ill,  .l/.J,  '..>VZ, 

\y\<i\:i\»-    Inr    iin:i-ui  in;;    i.~|iii:il" 

.liaii;;.',    ."■"! 
Apin'titi'   .iiiin'.    iintiiif   ..r.    I  III 
Ar.-,  rclli'X,  7s  I 
Aii,'iiKiv,.,    SI.  i;|il 
Ar-ininr,   tin".,   lilii,   i;l'7 
Ari.iiiatii-   Mil|.li:iti'<,   i;:!!' 
Anliytliinia  nl'  sinii<,  lilii'.,  J77 
Aitrrial   inrsslllr.    UJ 

Alt.  lii-;.  111 III.."    ill.   II'"* 

All.  ri..--.l.-l..-i^.    .Iiasl..li.'    |ir.'s-ili.- 
A-i'Miti.-   a.i.l,  t'.i'".,   lii'i' 

\~|.|.v\ia,   :;!  I 
Assilllilatinll    liinil.    t'.".J 
As-»...-iati..ii   ;ii.as,   ..i.'t.ial.   s;-:.',   sti 
iMiiit'ii-^,  7s;;,  7s.*. 

Aplasia,    .•.■lvl..'ll.-n-,    si  7 

Astli.'iiia.   Mi7 

\siliiiia,  .l.'a.l   -I'a. .     in,  :;l  1 

At.-iw,    .•.■i,l..'llai-.    si.i; 

\t.,ii'ia,  .•.■!.  I.,  liar,  si;7 

Al,.iiliaii,  ti'.l 

All-.. pill.',   ifl'.-.l    ..II   ;;laii.[-,    \1-1 

Aii.|it..ty    .-.lit.  r,    S.M 

Aiiii.li',   |.r.  -sill.'   in,   I  Is 

l.r..|.ai;.'ili.iii    ..]'    L.at    in,    IHI 

Aiiri.-nlar  .nrv.',  .■..iit..iii    ..!',    l'.". 

Auii.iihir    lil.lill.lti..n.    lln'.,    ->■'.<.   U's. 

Anii.iil.ar   lliilt.-r,    l:»i.   ->'<:<.  -''■' 

Anii.iil..v.'iitri.iilar    ..lili.-.,    I  Is 

l.nn.ll..   Is:; 
n. ,.!.•,  is:; 

\M-.,iihit..rv    in. 'til... I       ,.f    1,1.1..  I     |ii 

i:';ii 

Aiili..:italy-.i<,    77 

Ant. mi.'  11. U.S.  .■.r.'l.ral,    1.:; 

sMnpatli.ti.-,    i--. 
Ant. .11. .mi.-  ii.rv..ii-i  svst.m,   s77 

air.'ivnt    (il.crs    i.t'.    Ss.-, 


\  .it. .11. .11. 1.-    iii-i  \  ..lis    s\  st.n.      <  '..111  *.l 

aXMit   ifll.'x.'-.   in,  >is:; 

l.iill...-.a.i:il  i.iitlli.w.  SSL' 

1- i.-.ti.r   lil..is  i.f,  M7H 

tun. til. IIS  C.I',  SHI 

y..|iiTal    plan    ..I'   i-i.iislnnli.iii,   H7S 

parasyiiipalli.ti.',   Hsj 

tli.Ta.'i.i.liitiil.tir  i.iilll.nv,   ssii 
intrinal    M'si..-il    >pliiii.  Irf.    ss.' 
As. .11,   7sl 

ivll.x.  s,  7117,  ss:; 
A/.l.ii.-    a.i.l,    71J 


li 


ISa.illns    n.li    i-.inillllinis,    .'.IMI 
lia.liii.i,   ill    iiitistiiii',    lii'.t,   I'.'." 

in  stuTiia.'h,  4s:i 
Ita.tiTial    (lini"'li"ii.    •!''•' 
Halanic,  eiicryy,  .":!'i 

iiiiiti'iial,   'il:; 

.^liii.t   lit'  li.i.ly,  :.  1:; 
liaiilint.'   iiii'i',    '."1 
I'.asil  lirat   pr<i.lii.ti..ii.  ■":;>« 
I'.asal   rati.m,  ."i7r> 
lias.ipliil.'   .ells.  !Mi 
•■Hriiils"  III  ciiinsDii  wnrkii's,    iuu' 
l!i.|i/.iiic  a.i.l,  <i:ii>,  "111 
»fii/...yl    .-lilnii.li',   r>;'.i 
l!ciil.ri'i,    .".HI 

Iti'la  liv.ln.w    l.iityii.-   a.i.l,   "nil 
Mil..,  i\-Z 

an. I  fat  .liK'.sti.ni,  f.H'i 

ili.Miiistiy   c.f,    I'.il 

.■..iistilii.-iits  ..t'.    Wl 

fn.iii    yall    l.lti.l.l.T,    I'.'l.' 

fniiitii.ns   nf,   -lli:; 

|.i^iiii'iits  111".    I!'"' 

salts,    lilt 
liiliriiliiii,  4!'". 
Itilin-r.lin,   W'< 
r.  illii.laz..lvlrtlivl;illliln-,      .ir.'.-t      (.11      Mtin.l 

m'ss,.N',    :;..i7 
r.ii.ls,   r.Miinval   ..t'  Tu.-i    I'niiii,  HIS 
l;l.....|: 
■    ;il.s..rpli..|l    int. I,    I:; 

aniiiiii  a.-i.ls  in,  liml 
.  am. .lint    ill    lii..lv,    l:;."i 

antil'iTiiifiits  nf,  Sll 

.11. Illation    I.f,    I-- 

.lissii.iati..ii  iiir\r  nf,  ,';s:; 

fat    nf, 

.■stiiiiatii.il,   •'.!••!,  tiilT 

vaii.ati.iiis  ill.  IlliT 
f.iiinnts  nf,   H'l 
;;as.>s  (if,  Iraiispiiitatinii,  :;7;i 
y.'iii'ral   pn.pi'rti.s  ..f,  s.. 
iii.ass   in..\.'iin'nt    ..f,   l^sl 
in. ■alls   l.y   wlii.li  ;;:is.-  ai.>  .  .'ini.'.l,  :'.!lil 
i.xl.lati.iii  ill,  'lll'i 
pn.l.-as.^s  .,r.  S'l 
pr.itcilis    nf,    .s7 

i.ii^in,    ss 


i  I 


\A 


■A 


m. 


\ 


ss!l 


i;i I    (-MMi'.i 

i|U!iiit'ty  (if,  III  lupilv,  S' 

r.fiJIi  tivr    ilhl.A    of,'   NH 

s|ic(ili.'   nr.Tvity   (if,   sil 
Kiii:;ir  tcM'l  (if,  I'l.'iT 

rc;,'iilati<iii,  li'l 
triiiiifiixidii    (if,   I':!,    l.'."i,    l;','.l 

\iH((isil_v    (if,    I  Id 

Vdliiii f,   rili 

w.itcr  cdiiti'iit   iif,  ^li 

111 I   (■.■II,   red,   fiitc  (if,  !i:! 

(iriyiri    (if,   !IL' 
ic;;in('r;itliiii   (if,  !•". 
siKiiiia  (if    !l| 
white,  ii<; 

lllddil   cldttiii);.  '■>•< 
li>   (lisciiscs,    1 1 1 
ill   |i|i_vsidldKic  cdihlitidii^,   llil 
iiilliU'iu-c   (if   (-.'ilciiiiii   (III,    |i>.; 
iiilliifiii'C  (if  tissiii-*  tin.   Id  J 
iiitnivii-iciilar,   l".' 
iii(>tlid(N  (if  ict.'irilini;,  iti  ilr;i\vii  lildd'i, 

!Mt 
lu^atiw   |ih:isc  df,   InS 

tl ricx  (if,   Iliti 

time   (if,    1"0,   lOS 
visililc   (-lijini;('s  (luriii;;,   !m 
Ulddil  idrinisclcs  in  ludtilitaiii  sickness,   Idl 

It) Illdw: 

iliiiiiiil   cdiiilitioiis  alTeetiiiu  aiienii:),   2S,'! 
cTrtiidvasciilMr  disi^ascfl,  2*^1 
fever,    284 

(liseanes  df  iiervnus  system,   28-' 
mass  iiidvi-iiieiit  (if,  2'*X 
movement    in   veins,   21  I 
variatidiis   in,  2H2 
velocity  of,  2rt(; 
viscer.il,  212 
Hlodil   j;as   mandinetir.   .'ISI 
lildd.l   platelets,  i)7 
lllddil    pressure,   122 
diastolic,    127 

effect   of  lieniorrlia^'e  (in,   i:;."i 
e(T|ict  of  plenral  picssiive  (in.  .'Ifnl 
factors  maintainiii;,',    1.1 1 
11  idn   df  lildod   (111,  2:17 
mean  arterial,  12:i 
in  shock,  2!Hi 
systolic,   127 
traein;f,   12" 
lil.id.l    vessels,   ,8Sil 

el.-isticity    of,    1  12 
tone   of,   2.'lt> 
'■(idy  lliiids,  reactidiis  (if,  ll.l 
iddy    \veij;lit    and    emrfiy    piodiictidii,    ."i:i'.' 
lotiilism,  .'id.'l 
>d»nj.'in,  capsule   nf,  ."d7 
liadycardia,   llKi 
Irain : 
circulatidii   in,  217 
v.isomotor  nerves,  2.'2 
\ Illume   (if.    2.'iO 


llicatliiii;;,   in  ruinpiissr-l   :iii-.  illMI 

in   rari'lied  .'lir,   'Mh< 

perid(|ie,  .'lii:i,   :i71,   .'I7l! 
Itniwnian   miiXciiicnt,  cdlhiids.   .'7 
limit-.,    l.'iS 

Itiiffer   action   of   Id I.   .■;7  I 

llnfTer    siilist.-iiucs.    .Ill 
Kiiildin^  -tdiics   i,f    pnitiiii.  .',!i7 
linllidsaiT.-il   diitll.iw.   HS-J 
liiityric   •■1.  id.   7d:i 

(1 

•  'adaverine.  i'i21' 

•  'iifTeine,  li.'l.'i 
Caisson    disease,    |ii2 

cause    df,    lii.l 

decdinprcssidii   (if   uuikcis,    Iml 
;ire\('iitidii,    tm 
syniptiinis.    |ii2 
workiii;;    ciinditiiiiis    in.     Ins 
I'al.inni   idii,   inllin-iice  (in  iddttinj;.   In: 

inlliience  (in  heart,  Itil! 
I'alidnni    ri^or,    Kiti 
I'aldiiKl    (dcctrode,    :iii 
Calorie,   ."1:1.-1 
C.il.iriineter,  .".'l." 
,(iiitii;il,  ."111! 
llenidict,   .'1:17 
liomb.    -.:i7 
hand,  2H1 
respiration,  ."lilii 
Kiissel  Saye,  .-i:;7 
Cjilorinii'try.   dirci  t,   .-)l'i 

indirect,"  ."iHl,  .-.-( 
<*:inals,    semicircular.    H7.1 

reniin..|l    of.    S7 1 
<'aiitialiin.   .*i77 

Capill.-iiy  an.-ilysls  of  cdlldi.K.  •'i! 
C:irlijiiniiid   re.actidii,   ."i!'!' 
Carliiiliydrates,   alistirplion    id',   *l-">7 
••issimilatidii   limits,  <l."i2 

diyestidll    of,   (l.'ill 

,'iiid    ;;ni\itli,   ."is.'l 
UK'talidlisni   df.  Ii."i2 
I'riiductidii    fnim    pridcin,   liii." 
satiir.'itidu   limit,   ri,-2 
Carl  Kill   lialaiice,   .117 
C.irlidii   didxidc,   coniliiniii^   power.    12 
efTect   on   respiratory   center.   :i-"i2 
estimalidii  ill  lildod.  :iilO 
(intpul,    .■."id 

vcdnine   pcrccnta;;i.    in   Iilood,    '■':'.'\ 
C..||  l,,in    didxidc   tclisidll,   .'1.'17 

in    alveolar   air,    after   exercise, 
estimatiiin   of,   .'!:!',),   :t4  t 
in    mduutaiii    sickness,   .'Kil 
in    periddic    lireatliinc;,   :;7.-, 
in  arterial   Idood,   ,'!:17 
in    veiidiis    Iilddd,    ."I'.* 
larlidiiic  acid    '  .vi  /    Carlinii   dinvidei 
Carlidxyl   jirciup,   ','.>» 
Cardiac    decdm]iciisatidn.    Ill  I 


I 
III 


S<t(» 


iM>i:,\ 


,1    ' 


Wn^  i 


1 
j 

! 
i 

C'lnli.ii'   ilipn xHiir    mrvr,   LM!' 

I'lircliiii-   mil-"  If.   |ilivsiiilii){i.'  cliarin  Irrislim 

.if,    i:ii 
f'liriliiii'    iMPihli    I  itiiiMiuli  I,    I'll 
Curiliiii'    »|.|iiiirlrr,    lis 
('iinlii>rihal  ili><i'iis.>,  liliii..llliiw   in,  ;;"<l 
riicrnv  (iiit|iiit   ill,  -ilL' 


('.•iriliciyriiiiis,   -I 

I'lircliiiviiHiuliir  i|i«i:i»i',   lilcMi'ltlnw    in, 

CiiM.'in.   IMS,  ."iTil 

I'lisciiiiipii,    ISS 

I'litalasi',  !»iP 

ration?*,   lit 

<'atal,VMt'<,  7l' 

•  'iifiilvtii'  pdHcr,  LM 

Cilriitrratis.  iirrvipin  HVstfni   nf.   ''*- 

I'l'lliiliisc,   I'ltirstiiiii   of,   .'iiHi 

I'liiti'is: 

iiKsot-iatiitn,   h.'iL*,   h.Vi 

ilialn'tif,   tlTl' 

motor,   HV'i 

aiiilitor,v,  H.'il 
visual,  H.*)l 

xcnxory,    H."iii 

Word  "<ciitcr-i.   sCii' 
Ci'lilialiii,  tlsii 
I'.i.alH  aihl  yronth,  ."iHI 
C.TilM'llar  ataxy,  Hiiti 
( 'rri-lirlllllll  ; 

alilatioii  of,  SfiH 

rliiiii'al   ohsi'rvatioiii,  xT" 

fxtir|iati   M  of,  silll 

fiimtion       .f,  HC.-, 

loli.s  of.   Htls 

locali/.atioii  of  fiiiirtioii  of,  HiiT 
f'i'ii'tiral  cinulatioii.  JIT 
I'rri'linil  riiiii|irisHioii,  'J,':! 
CiTvliral  rortcx,  Ntiiiiiil.-ition   of.   SH 

struitiiri'   of.   H,")L' 
Crri'liral   loi-alizatioii,   si:i 

c-liiiiial    olisirvatioiis.    N4!l 

lii'nu'i|iliiTt'>'.   n'liioval  of,   xi" 
I'l'ii'liral    vi'Hsrls.    ligation   of,   -I" 
(■(■n.lirosi.iiial    lliiiil.   '.MS 
('rri'liniiii,   liiylirr   fiiiictioii«   of,  Hiio 
<'ll,  inc'tlioil  of  cxprrs-iiiK,  -7 
I'll. ■VIM' Stokes   lircatliih),',   :i71,   :!77 
Chloridrs,  ii.iiii',  ."i:i 
Cl.ol.-MiTol,   4it4,   (iS.S 

I'^tiiiiatioii  of,  <I!i7 
riiolinc.  tiHjt 

Cliorda   t.Mn|iaiii.   'J:!!,  :;ilii.    I-':'. 
llir(.inatiilysix.   MH 
Cliroinatiiu'.    •'•IS 
Cliroinosoni's,  ti:iS 
•  •Iiymi\  41<!,   4H2 
Circle  of  Willis.  1117 
I'irriilation  of  liloo.l; 

lontrol   of.   ;;i() 

iiilliu'iicc  of  jtravity  on,  L'44 

mass  movemi'nt  of  hlooil,  lii'H 

tlironuli  till'  licart,  L'.'i7 


.,^^ 


firiMilatioii  of  lilooil     Conl  M 
lliroiiuli   till-  livir,  -"i.'i 
tllrllll^ll  till'  liiiiK",  -'>'l 
tiiiii'  of,  -.'i:! 
(in  Illation  tiiiii',  L'litl 

I'liiiiral   applicati tinulation.   -.'H 

nirvoiis   xyst.ni.   x:;S.   HIH,   HiiJ 
ris|piratioii.   :ilii,   :i'Mt 
riottinu  of   liloo.l    i><>    illo.>.|  I'lottiiiul 
< 'oai;iilati\i'    f.Tin.'iits,   XL' 
C.i.lliv.T  oil.   nntrilivi'  valiii',  7i"i 
•  •o.  Illii.iit   of  oxi.lati.in,  :i!».l 
Cm'rti.i.nt  of  H.iliil.ility  of  K""''".  •'•'' 
Colli   s|i..ts,    7!>- 
Ciillat.rals.   7SI 
Coll.ii.ls: 

llr.i»iiian    mov.incnt.    "7 
.'a|iillary    analysis,    .'i. 
.Iiara.t.'risti.i   |iro|.(rti.is   ..f.   "il 
.hfl'iisiliility   of,  ."il 
.lisptrsi.in  mi<niis,  .">4 
.li>|..'rsoi.l,  .*>4 
ili'.trii'  |iro|>i'rti.'s  of,  ■".•". 

osnioti.'    |iri'ssiir.',    .'.7 
i'l.'.tro|ili.irr.siH,   ."id 
.xt.'rnal  |iliasi',  ."4 
yilatinization,  ill 

li.'t.r.i^.'i us,  ."il 

lionioy.'ii.'ous,   .'I 

iniliiliiti.in,  l>2 

int.'rnal  pliasf,  ■'  I 

isiirl.ii-iiir    point,  til 

ly.ipli.ilii>,    nil 

iiiiitiial    pir.ipitati.iii    .if   I'.ilUii.ls,    ."lO 

osin.it i.-    pri'ssnr.'  of.    111 

si/..'  .if  i-.illoi.l  partiilcs,  53 

siisp.'iisi.nis,   .'i.'l 

siispi'iis.iiils     ami     .'iniils.iiils,     a.ti.iii     iif 

I'lortrolyt.s   on,    lil 
Tymlall   pin ii.nn. '11011,   .'.I 
C.iinp.-nsatril   a.*i.|.is*'s,   .'lit 
C.inipl.'in.'iital   air,  .'IIMI 
C.inipr.'ss.'il   air  si.*kii»'ss.   ;{1*!* 

laiisi'    of   syinpfoins,   4ii:i 
privcntion    .if,   4iH 
tr.atm.Mit    of,    4IM! 
C.in.-.'iitration  .'.•11,  :'.ii 
Coni'i'iitration   p.iiiit,  aiiri.'li'S,   lH."i 
Coni-fptiiin,    Mlil 
Coni-.pt,  .S(il 

C liti.iiH'il    r.'tlfxiis,   481,   H."i(! 

C.iinluitivity,   .li'trrmination    of,    17 
.'.{iiival.'iit,    1!' 
m.il.<.'iilar.   !!• 
sp.MMli.'.    17 
imlintivity   .'cll,   IH 
iii);liitin,  ."i.iS 
'onstru.-ti.iM    .if    aiiton.imif    ni'rvous    sys- 

fi'ni,  877 
'.mtra.turo,  fxtcnsioii,   800 

iklll^,    oW.'t 

'oronary    i-iri'ulatioii,    -ol 

'oronary   vi'ssols,  vasomotor  norvi's,  268 


INDKX 


H!H 


68 


('iir|ii>rti  '|iiii<lrl);i'iiiiiin,  **(ii 
Hi'i'tiiiii  lii'liiiiil,  HJii 

Bi'rlliill    ill    friillt    lit.   Hid 

•  'oriniMcli'H  cif  lilipiiil,  ml,  Itl 

whitv,   llti 
t'lirtcx,  ri'iiiiiviil  of,  Hill 
<'i>iiKliinu,  ;i(Mi,  ^]•^ 
<'r!iiiiiil  ciivity,  iircHsiiri'  in,  'J."it 

•  'riiitliii',   <"»!,   tli:i 

rliiiiiistry  (if,  i'iT2 

otiiiiiitiiiii  iif,  •iL'l 

in  clisiiisi',  ilL'il 

iiii-tahiiliniii  of,  <'r.>l 

nrlKlii  (if,  'ilMi 
I'ri'iitiniiii',  liKI 

iliiMiiistry  (if,   tlL'J 

(11.  ftiiiciit,   tiJt 

istiiimtidii,  t'lL','! 

ill    urine,   '>'M 

iiictnlKilisiii,  ti;} 

(if  hI(Mii|  ill  (|i>4i-,'iHt',  fi.'l 

(iriKi'i  (if,  ti-ii 
''ii»la  iiinxliiii,  ST'.i 

•  'ritiiiil   ('iiiici'iitrntiiiii,   H 
t'riwscil  cxtciisiiin   rcllix.  Hill 
Ciiririii,  tiHIt 

•  'iiirciit  (if  iictiiin  (if  licart,  1h7 

<'V!lllllsis,    .'ilill,    4011 

•  'yKlciiic,  liii.'l 
Cystine,  .'i77,  ."iltL",  t'liil 
I'yMtdsine,  ti:!7 

•  'vtascM,   4<i:\ 


D 


Halinatiaii  iln);,  |iiiriiie  inetalMilisiii  (if,  till! 

I>all(m  's  law,  :i:iii 

Keail    «|iaii',   mil.',   .llH 

Ileafness,  Htil 

Heaniiili/atiiiii,  ileaiiiini/atimi,  .'111 

l>eaiiiiiii/iii^'  eii/yiie,   ti:i!t 

Decerelirate    rit;iility,   HUH 

Decerelirafidii,  HII!  ' 

l>ei(il(irizatiiiii    nf    liijiiiils    liy    .  Iiariiial,    lit! 

neidiniiressidii,    IIMl 

defecation,  47ii 

lilddil   pres-iiire  iliiiin>r.    IIL' 
Defilirinateil    lildci.l,    Inl 
l)et;eiieralidii,    siicce-ssive.    Hi:; 
I)et,'lulitidii,  44"i 
Melayeil    i-diiiliietidii,    l'7ii,    '27t> 
Oeliritini  eiirilis.   111."! 
Dendrites,   7H4 
MepresMidii  (if  freeziiiy  |idint,   li> 

(if  urine,  .'I'J.'i 
I)e|iie!isiir  nerve.  I'HH,  LM!*,  LMn 
Ileiiresudr    suhntaiiees,   .'lil7 
Dessert,  iihysidlojjii'  value  df,  4,'!7 
Detoxicatioii  eoni|idunils,  6'2'.l 
Detoxieatiori   jiroii'ss,    "iiil 
Dextriiis,  4!M,  «,-(> 
Dextrose  (are  Glucose) 


Djalieti's: 

aci'limii  in,  liHl 

nil. I   the  (luctl.Hs   ijlaii.N,  r.7H 

n^Kiiiiilatiiin  limits  in,  il'iJ 

lildiiij   exaniiiiatiiiii    in,   (i.'ili 

IiIoihI   fat   ill,  iMi'.l 

(I  liter,  'lialii'lic,  t'i7L' 

early   dia^fiKiM"*   tif,  tl'ij 

eiierKy   (iiit|Mit    in,  ."ilj 

e\|M'riiiieiital,  *i7J 

fat   nietalidlisin  in,  )iH:i 

ketiisis,   tiH:i 

paiiereatic,  (17H 

iier\'dns,  in   man,  ii7l 

|ieriniineiit,  li7t! 

phldrlil/in,  (Hi'i 

|"i-.l|iraiii|ial    liypitul\  .iiiiia.    ti'!' 

renal,  li'il 

xtarvatidii  treatiinnt    in,  (in  I 

treatiniiit   df,  ti.'i.l 
Diiilietic  aci.liiKis,  liHI 
Dialictic  center,  t'i7:; 
Dlalietic  ;{aiii{rene,  li'S 
Dialiirii    .■ni.l,  lil". 
Dialysate,  ."J 
Dialysis,    IJ 

inetlliiil,    ciilliiiils,    .'il 
Diaplirayii.  actiim  nf,  .'Il'h,  :'i1!I 

pliy«idliii;y    of,   ll'.'l 
Diasfdiii-   liiliin;   ,,f   heart,    l."i.l 
Diastdlic   pressure.   lL'7,    l:il.' 

ineasiireiiieiit   (if,   in   man.   1-H 
Diiriitic  iidtch,  L'ii2 

wave,  L'ii:t 
Diet   .at   (lifTerent  at;es,  ."i!MI 

of  ilill'erent   cdinmiinities.  ."iH!> 
Dietetics.   ."HH 
DitTercntial   iinncurietei .   .IH] 

DifTiisi 1:; 

|)ii;estilii|ity    df    fcdils,   r.<t:i 
Di;;estidn.  liy  paiicicatic    juice.    |H!I 

ill    iiilcstiTie.    |H!( 

in  stdinacli,    IHl 

II Iiaiiisin  (if,    1 11 

Diye~ti»e  ylaihls: 
cdiitnil    df. 

Iicirii 4'_'"i 

iiirvdii,«.    ll';t 
j,'eiieral    pliysi.il,i(;y   df.  41 S 
micriisi  iipic    cliaiiyes    iliirinn    aclivitv, 
4IH 
Dispersion  iiieiliiini.  idllniils.  .'1 

Dispersdiil Iliii.ls.  .",4 

Dissociation,   Hi.   17 
Dissociation  constant.    l!t,   :1HS 

DisS.lci.-ltidll      curve; 

of  liloo.l,  .•tH:i 

of   lienio);loliiii.   .'Ih;; 

inlluence  of  salts  on.  ;(H.'i 

iiithieiHe  of  Hioii  ooiicentration  on, 

:i8(i 
intltience  of  foniperatiiie  on.  :iHti 
Dissociation  liypothesis.  applications  of,  21 
Kissociation.   rate  of.  .'tHii 


u 


'  ■ 

ill' 

'  1 

III 

i\i, 

iMur.'li. -.,  .■;'* 
I'lM  r  'x  piil'V,  lti'.' 

I tihlH    lllrilli.<l.    '.I  I.    '■'* 

iir,.|.|u.i  i,.iii.  ■.■:i 

I  III   Iti.U  ruriiiulii.   '>ll 
|iii.  Ill— I  uliin.l-.  T-!' 
ih    .li:il..ti>.   r.> 
l>>»|.Miii.  :;i  I.  ::i!i 

|l\«tf.i|.ll\  ,     r-..l;lti-.ll,     X"* 

y. 

Kiiilli    WHiiii.    hiU'iiix    ,i.tiiii    I.I',    7>>il 

i:.^  li^iiii:!,  i;i7 
i:.  i,'iiii{'>iii.  I'c'o 

IMiiiia.  t'li'.   IJH 

Ivli-tiii    iiii'l    uiinith,    .''^T 

Kla»liii.  .|iy.'-li..ii  lil',    I".!'. 

Kh'itiii-   .  i.hiliii  tn  it\.    111 

Kli'i'lrii'  i-iirri'iit«,  iliu'lupiniiil   i>t'.  •'.' 

Kli'i-lrii'  iiri.|MTtii"4  lii'  ihMmM-.  ."..', 

KlirtriHar.lii.i,'i:iiiii.    1."s,  j.-,:i 

iiiir:  .il,  :j(ll 

^l:iii.|ar.|i/,:iti I'.   L'lln 

viiitrii  iilar    i  ii1ii|i1ia.    u'I'- 

wavt's  (if,  *2*i\ 

r  «aM'.    I'^'.'i   -111 
T  »aM',  -Jii.  '.''i:'. 
Kill  tin.  :ir.|i.i;:ra|>)i,   '-'I'l'i 
Kli'i'triii-iitiiiii.  laii-i'  i.f  il.'aMi   in.    IJi'i 
i:ii'iri(.IUi>,    lii 

ailii'll    llf,    (HI    Clllni.l^,    I'l^l 

Kli'i-tni|\tii'   siilutiiiii    |irrsMii'i'.   I'l' 
Klc.-tr(.|'.li(iic»ls    I.f   i-iilloi.h.    r.i\ 
Klcitrii«lalir    attraitiiiii,   -.".i 
Kiiil.i.li.   in: 

i-:iiictii<,  i.-iii 

Kiiii  tiiinal   nhcii^iiiia,  liT" 
l'.rn|.li,\>riiia.  ':!ll,  .il  I.  :;-l 
l''.in|i\iiiia,   ''ilM 
laiiiiM.iii--.  iWs 
Kmiils.iM-.   ii.lli.iiN.   I'm 

llinliiri iiii'  iiiiiaii'i.  7i;:i 

l-]iiiliiiti/\ mr.   71 

i;iii|ii-illnll-i    llirtal"ili-lll.    I'll.'. 

Ill'  I'liiims.  tit" 
Km  i-.v   lialancc.   .'nr. 
Kiii'iyy  iiiitpiit,  ami   .-iLir,   •'  1 1 
.•ii'iil   limly  w(>i;;lit.   .'Ill 
ami   iliM'!!"!',   •'!- 
ami    inusiiitar   wmk,   ."  I'.i 
ami   -ix.   ."ill 
ami   ■■nrl'.-ii-i'  .an  :i.   '■  I" 
in   slaivatinii.  ."IN 
IjitiiiikinaM',    tl'l.    !><:• 
Kii/viiii's,  71 

aitiiiii   llf   triii|ii  ratiiii    ini,   7  I 
aaivhiM-H,    H\ 
ami    fatalv^i^.    7l' 

aillii'M/\  llir^.    '^l 

.•ii"iii.i~'.'.    -1 


IMH  \ 


Itl/llili-       I   i.hl  M 

■  n.a;:iilaliti'   fmnrpil-i,  l«'.' 

IlliliMI    (if    iiithitv,    I*'.' 

I  (i.|ii(n/_\  iiii-M,   7! 
ui\  ii\\  laic.    XJ 
iincMaii-.  il 

I i|.n •**'•*,    ^1 
naliiir  "f,   7- 
nxiilasi  ■*.  ^- 
I'l'i  iill.-irilii-i   llf,   >"i 
I'liiixiiJascK,    ••:.' 
|iiiiliiftic»  (if,   7-1 

piiitraiii.    Ml 

II  \iTiiliility  I'f  artiiiii  I'f,  ^'i,  77 
-I'ciilir   Ilitiiill    llf,    7  '• 

l>|ii'i  llf,  7!» 
iiiciisi>,  H;.' 

»rl(liit.V    liillililllt,    71 

K|iiiritli'   rci  r|i|iii'i.  7IIII 
K|iili|i»y,   .lailKiiiiiaii,    slit 
Ki'ini-|i|iriiit',  'J.'lli,  .'h- 

:iml  ili;il"t("i,  117 :'• 
Ki|iiillliriiini,   iiiliii;,'i'ii,   .~i71 
Ki|iiivali'iit,   iiimliiiliv  jly,    11' 
Kiv|  .ill.    Mill,  fiiil 
Kruastiiiilami,    I'.'n 

i:r«i,t,  r,(i-2 

i;itfiitii\iiic,   'Jii!* 
Krvtlirocjli'x,  :•! 

i'atc  llf,  !•! 

i(';;('iii'ralliiii   iif.   'X\ 
INialMiiii'iit,  L'lH 
Ksii|iliayiis,   iliirim;   sivallnu  in;;.    I  I'i 

inliiliitiiiii  of,  417 

IK'iixt.'iltic  wave  in,    1    7 
IMirs.  liHO 
Kstrr  viilup.  ti.'<7 
Ktlicrral   siilpliati'.K,   ."iiil,   H:'.- 
Kxi'i'lsin,  ."77 

Kmuji'iiiius   iiirtaliiili.sin,   fil  " 
K\ii|ilitlialiiiii'   t'liitiT,   7."i<i 

cncryy  (iiit|iiit    in,  ."IL! 
K\.  ii'tiiiii  llf  aiiil  iiiinliiniil   rtilli  aiiiimiiiiil 

ti'i 
I'.Miitiiin   llf   uiino,  ."iC 
Kxti'iisiiih    iniitnii'tiiri'.    I".    ><«' 
Kxtciisiir  tlinist,  ."i7 

nllcx,  Hitr, 
K.\lciiiiT|ifi,rs.  7MS.  si;'_' 
i;.\tr:i^v>tiili'.   lii'ii'i 
|:m  -.    I'liiixniiilit-    llf.    si7 


Faitiir  safety,  ill  i1ii-t,  .''.'- 
l'.ati;;iii'  llf  rcllrxi-s,  Sij.-. 
I'at-: 

:ili.i.i|.li r.  li'.il 

rlii-inir;il  tliiiiiy.  •''.•:; 
ini'i-liniii'^iii'  tlii'iii  V,  liliL' 

t   ^jr.iwtli,  r,H^ 

l.liiiul,  lilMi,  (107 

ilistiii.atiiiii   llf.  I'lMii 


Ml:! 


I'll-,   IJ I     r,.iii  M 

i|iliriiiiii;itiiin,   tltii! 

i|iiiiii;j    iilisiir|'li"ii,   il'.'H 

■  liiiiiii;   I'a'tinj;,  i'i''H 

»;iriiiti      ■<   ill,   (;!>r 
I  lirllli'.tri     iif,    ilHd 
•lr|'ot    flit,    tl!i!>,    7"ll 

<N«tiiiiitii>ii   III",   Ti'l 
il> -11111111111111  iif,   7""i.    7IJ 

.liUI'tllMII    of,    tllHI 
flit    illlst,    li'.H! 

IImt  fill,  lil'lt,  Till 
iiii'IiiIioIhiii   of,   II'MI 
ti-«iii'   fill,  I'llHt,   'iitl 
triiiiviioitiitioii   to   livii,   7"^ 
l'iitl>    iiiiili,  i;sii 

ih-i'l   iiiiiiihrr,  rt'^7 

lircvikilowii   of,   7ii!i 

rstir   \nliii',   l'i'<7 

fiitiniilioti     from     mi  K.ilii  .|r:ilis,     ; 

in    liv.r   ill   i|i«>;i»r,   7"'; 

in. Mill'    Mlllll',    l!*"! 
Illc-llill;;     |ioil|t,     r.S7 
IJriili.Tt   .\lfis,|     Mil,,,,     liss 

Mi|M,iiiiii-'  lion    Mlllll',    os7 
I'l'. .-,    I!t|>,  ,,-JI 
ri'iiiii'iit-i    '  «,.     Mii^v  nil- 

h'l'I'Mlflll'i     ill     1,1, Hill,     Kit 

I'i'Mr.   1,1 llloH    ill.   L'xl 

fol.l  l,;illi   tri'iitiii.nl.   •Jsl 

imriiii'   i-\,rrlioii    ,liiiiii|;,   I'.IS 
I'iitrr-*.  iiiitt'i'ior  r,,ot,    Ion 

ii>llllrrti,r,    I!'.'! 

iiiti'riinii,  iiil.    SiiJ 
I'ilirilliition,  iiiiii.iiliu,    I'.ir,,   i'i;:i 

•  rnlriiiiliir,   l!'."i 
I'il.rin,  '.I'.l 

(iliriii   iiiM',lli>.   :>!' 

-.oiiiii'   i,f.    Iiil 
I'll, rill   fi niii'iit    I  M  .    'riii,,nilMii    .    In;.' 
I'il.iinok'rn.  ^7.  l"l.    I'l;;,   111 
I'ilniitioii,   1.1 

I'iiiiil   I'oiiiiiioii    |,:i'li.   7'<7.   *>.'l.   '•J  I 
I'istiilii.   Iiiliiirv,    I'.i-' 

t'li-tii,',   1:11' 

sjiliviirv,    t:!i> 
KliAioii  rrlli'x,   Mil.    Hu'l 
I'liittiT,  iiuriiMiliir,  :Jil'.i 
roo,l : 

ai'ri'ssiiry  faitor^  i,f.  ."i|i:; 

I'liokiii;;,    iiii|,ort,'iiiri'    of,    ."i!i;i 

fITpi't  of,  on  i'iiriiliitii,ii.  L'l;; 

I'lTfi't  on  en  iitiiiiii''  i\,T,ti..ii,  liL'l 

lllMllivi'    l|lllllitii-.    .",|i| 

piil.-italiility,  .'I'.i:; 

r I   faitorsi,  ai,  ,■>-,,,  V.  -.sl 

I' l.-tiifTx,  mil'  of  l.iiMii;:   -Liinaili,    i:.> 

riirrcj   lirpatliiin;,   'i-'l 
Koriiial,|i'liy,!i'    titialinM.   ;iii,iii,,    mi, Is,    J 
Kori'iati"!!    of    s,,!i,l    siirf:,.-.-    '.:'.!<::,    ill! 
l-'ri'i-zinj;   point   lonsliinf.    In 
Kriiv.iii;;  point.  .|i'i,n>!ii,,ii  of.    in 


"I. 


I'l  i.|,i  I.  Ill '-    III,  111. m1    f.,r   iihi.iliir    air,   .'II 
rriiiiliil   \i>.iiiil  .inter,   s",| 
I'mi  l.i«r.   tiiKi 

riinrti,,|l<     of     l,,ll,il|,,||lii       iH'ltia.     ss| 

r,i„.|ii«  ,.f  «lo„i:„  h,    |-,i' 


a 


lliilM I!l| 

lialMiiionii't,  r.    ^lll!'-,    Is7,   L''.M 

"iiini.'iiii,   7><l 

liiM   ill  •itoniii.  I,.    H'lJ 

( iiiH  liiH^,  ;:,  ;:,:ii 

•  iiixi -,  H.l-orplii.ii  ..I',  111'. 

roi'llli  ii.iit    of    „,l,il.iliiv.    .;  ;7 
i"-tiniatioii  i.f,    ;  1 1 
partial   pri«>iir.'  of,   .;.;i; 

solution  of,  ;i:;it 

liii-ion   of,  I'.'ti'i 

lriiii.sp,,rlii;ion   in    1,1..... I.   :;!iii 
iiaski'll's   riaiiip,    17". 
Hariri.-    i-onti  Ills,    1.  L;.n;;ili,li f,     I  111 

l,||s|li.-      .liliOSti (»| 

rat.-   of.    Ih7 

•  iastrii'  fistula,   I.U 

fiastrif    jiiiii'.  i|iiaiilil\    -.ri-ri  li'.l.    I  pi 
striMisitli    of.     Ill' 

liastrii-    ..■.T.li |:;l' 

liiiriii,,ti,.   iiiiiirol    of,    .i::7 
loi-al  stiiniilatioii  of.    i:.s 
iiirvi.iis   ii.iitrol   ,,f,    |:;| 
liiistri.-    tiilii.,    I.V; 
I  last  rill.     I.lli,     ir.ii 
liastroriilirosloiiiy,     lil.i 

liristrointrstiiial   I'oiil.nts,    iii,.ti,,ii    ,,f.   .'.ii.', 
<i, latin,    :,7s 
'"■liiliiii/alioii,  I'.l 
lilaii.ls,    .liMliyrs    , lining    a.livity.     I:.'J 

I'lirtrii'   rlianoi's,    Il.'J 

ii.,riiial    roinlitioiis    ,.f   a.ti\it\,     l:;ii 

oxygon    niiisiiiiipl  ii.ii    of,    :.|ii;.     |JI 

II  Kpiration  of,  ::'.'i\ 
liloliiilin,    .".77 
I  ilia. Iiil,   .-71) 
I  ilonii  riihis,    :..i7 
i.liiii.ir'oy.iii'.is.   liili'.    r,77,    i;s,, 

•  liri'i-t    ini-tlio.l.   i;r,:; 

iii,liiTi-t     inotli,,.|.     iir.l 

in   iiornial   aniinals.   r.ri7 
t'ilii.  IIS,..    7im 

flit.-   ..f   iili-..rli..,l.   i;i;j 

i;lii,o»i.   to   iiitiiiyi'ii    ratio,   I'll!  I 

ill  ii'i-li,,tis.    iiitiai  ,t,,,iis,    1;.",:; 
sill, .iilaiH  ..lis.    lir.r, 

paii'iitrial    assiniiliit  i..,,,    ii.'t; 

l.iji'ran.i'   f.ir.   il".  ; 

iilili/iilion  of.  in   li^xii..,  ri77 
til.ilaiiiii'  ai'i.l    1  M  .    i;iiitaiiiiiiii'  ai-i.l  1 

•  iliitainiiiii.   aiiil,    fio-'i,   rii;7 
(ihiloiii,    ,",77 

i.iiit.  iiii,  .'.77 

(ilVirir   al.Nl.vl,..   iHl.-, 

(.l\.-,.rol.  lii'io 


t  .t 


8!t4 


IMIl  \ 


\}'.i  i 

::i-:!|i 


!  '^ 


! 


lifllCMS  ) 


(ilvr(MlM>li<'  :i<i'l.    I!M,  i\:\] 
(;l'vrinr,    I'.M.  lin:; 
(ilyciiiin,    .'77 

»;iyiMiiM)ll,  till],  ''>:;ii.  iii''7.  71ii 
<;lyi'<i;;i'n,   •>(!'_' 

Villi'    (if,   t'lCi.t 

Miiiri'ii    iif.    I'l'l- 
(il\iiii;iMi:i-i'.  I'li'iL' 
(llyi-.i;;,ii..l\sis.    (;r,l> 

liiiniiiiiii!    i'i7ri 

llrrMill-.     *U~ 

jiii-itiniiili'iM.  i'i7ii 
ClyidlMl.li'hy.li'.  ti"-|.-. 
(;lyi-iily^is,   ri77 

(ilyi'iiiiiMiyi'iirsii    I.V.I     lilui- 

(ilyi-iiMiri.i,  iiliimiit.'iry.  I'l.'ili 

IMIIIitiiill.'ll.     t'i7"i 

|iiist]ii;iiiili:il,    *i"i'.* 

ri'hiliiiti  111  Mi^Mi-  III'   III I,  "i'iii 

rriiMJ.  i;iil 
<ilyi-iir(in;iti-^.  *i-"'|' 
(Mviiiriinii-   :iiiil.    li^'iH,   'i-l.   '''■'•- 
<ilyiix:il,  ii:'.l 
(ilyiixylii^i'.    >>-.    lii'ii'i 
(llyiisylii'    mill,   I'l'.l 
i.  N  rill  ill.    tii;i 
(iiiilrr.  i-\ii|ililh.'ilinii  .    "i  IJ 
(mini.    I'll!*,    ri.'ii. 

.■tiiili.;;y  111',  I'l-".!! 

^naiiini',  'iP' 

iirii'    :ii-iil      siTi'tiiiii    ill,   lil** 
(irii.lill^:   III'    iiili-n-ity    i.l'    nlliA    iirliiiii.    '^'i'.> 
(ir;nu    !niili'i.r;i',    il,    'i 
(li-.'itii    iiiiilrriihir    siiliitiiih,    -•-' 
(inivily.    III!    iiiiul:itiiiM,    Jll 

i-tirinii'li^:iliiiM     I'lir,    "J  t  "i 
(IriiHlli,   "71 

.■11  T.-isiiry    I'.-n-liir-i,    •"'v'l 

ii,*i-,;(I    ?-,*ilitiri,  ."i7*i 

i:irl.ii|iyilr;itr.i   an. I,   .'|^:: 

.iiivis'nf,    ."7i; 

1-nrvr.i  111'  iiiliiliiliiiii,  ."i7!' 

t'.'ili  iniil,  '•<■ 

iiiiiru'inii'  valN   an. I,    ".Mi 

lysinr  ami,   •"i7'» 

|irotrins  ;mi.|,   ."i"  1 

liyiianii|ili;iiii'   ami,    ."i7>< 

vitaininos,    ."iSt 
l.iianiilini',   ti""i.    li-'- 
(iiianini',  I'^l'i 

< iuanii..ii!!r,  t'l-IH,  *i;;i» 
(iiirisln'r);  n'am'iit,  4S7 


H 


ll.-il.l.'ini'  Haniiil't    a|i|..ar.iln-.    I". 
Il.i|.|:iiii'  t;as  a|i|iaralns,  ."i-'i!l 
llal'lam's  rnilliin'   fur  alvi'filar  air.  lU'i 
lliart: 

ai-tii f,    1  It 

aiiriiMilar    riirvi',    1":' 

iliastiili'  iif,    1  I'l 


lli'ail      Ciinl'il 
isiiinclrii'   |n'riiiil    in,   M!> 
iniKili',    |irii|irrtirs,    17i! 
nutritinii     nf.   l'''l 

ii|ii.aiai:   a.,  I   i-liisinK   cif   valvi's,    l.'il 
ii\yi'  a    rriinimiiriils   of,   il'.Mi 
ii\y;;rn    >in|i|ily   nf.    liit 
|.irfn>iiin   uf  mitsiili.   Iimly,    liil 
|..,.iti|iliy;;inii-    |ii.riii.i,    l.'in 
jiri'^liliytfinii.    piTiml,    1  I',' 
jiri'ssiirr    in,    1  Itl 
{i>iin|iinu  ailiiiii   nf,    l-'  1,111 
ri'..iisrilal  ii"     I'l   stfu.    Itil 
rhvlliniii.    piAH'r    in,    I7ii,    171 
M.iia.l^   nf.    I. '.7 
systiili.    nf,    1  I". 

ntilizatiiin    nf    ylmiiM-    in,   liHl 
vayiii   inntriil    nf,   inM    lilniiili'il.    '-'17 
va;.'iis  innlriil    nf,    iiianirnaliaii,   --'i 
vau'iis    tcriiiiiiatinMS    in,    --'i 
vi.atrii'Mlar  riirvi',   1  Ki 
u.irk  nf,  '-Ml' 
1'   arl    lii'al: 

.'irrlivlliniia    nf,   -li'^ 
niyii(,'rni.-    Iiy|iii|ln'>i<   i.:'     171 
ni'iirn;,'raii.    liyimtlii'.ii.*    nf,    1711,    17'.' 
nriyill    if,    ill    rnlil  lilniiilnil    aiiiiiiaN,    1711 
<iri(iiii  nf,   in   iiiaiiiiiialiaii,    inj,    l>«!t 
]ia.i'  niakiT  nf,   171 
|irii|ias,'atiiiii    nf.    --' 
syMi|iatlii.tii-    (■nntrnl    nf,   -'-:i,    '-'-7 
ultiiiiitm     imn'u  ns,     1  H.'i 
va^'iK   inntriil   nf,  '-'17,  :.':.'ii 
II, art   iiiiii-k.    17t.  L'7ii.  L'7ti 

nfl'i'i-t    nf    vayiis   nil,   "Jll) 
ll.;irt    ilisi-.MM',   vilal   ra|iaiitv    nf   liiiiys   in, 

:',11 
lliarl  Itinu'    |iri.|iar;itiiiii,    I"'* 
llial    |iriii|n.-tiiiii    ami    au'i'   .■iml    >*n^.   ■"' " 
ami    linily    Hii;;lil,    'i-:! 
siirfai-i',   ."ilii 
ilisnasr,   .-ill" 
lliat  H|ii>ts,  7!'"-' 
Ili'.'it   valiii'  nf   fii(ii!>,  •"^'■•"i 
lli'iii.atiiiTit,  7 
i      llriii;itn|iiir|iliyriii,    l!''i 
'     Ili'iiiilil.'Cia,   '.'.'iS 

lli.|iiiiilr(ii«iiH;ra|ili,    -'"i 
llnninulnliin,    !>1 

ilisiiiii'iatiiiii   iiiiistiiiit,  .l"*"* 
ilissiii-iation,    i-iirvn    nf,    HMfi.    IIS-.',    ■.W:, 
nstiniation    nf,   !•- 
I         r.'iti'  fif  iiissin-i,'iliiin,  ^'.HtJ 
'  ri'laliiiiislii|i   In    liiln    |ii>;iiii'iits,    1!m; 

><|iii-ilii-   oxynon  ••.■i|iai'ity   nf,   :17!I 
trans|inrt.iiinn    nf    <)_,    liy,    .'tl'f' 
lli'innly^is,   7,  l'"i 
lli'iiinlytii'    janinliii',  !'•! 
Ili'niii|iliilin,   11- 

Ilnmnpoiptir  arlivitics  nf  liniic  marrnw,  O.'l 
llrninrrliatif',    ."i!* 

iiiimoiliati'  I'fTnrls  of,  1:17 
ri'i'dvcrv    frniii,    KSH 


INKIX 


89.') 


1711 


llriiKirrliaKii'   diMiisi's,    111' 

IIi'iilc,  loiip  ipf,  .'ill" 

llrpatir  artiTV,   llow   in,   -'>'> 

Ili'tiTdi'Vilii'   i'<iiii|Miiiiiil>.   Hill 

IlrXdsi'?*,  ti.'ii' 

H  ilirriiatiii^    niiiitijil,    ini'talmli'iiM    of 

llilii'riiatiiiii,    lii'i'.'itliiii);   iliiriiit;,   ;I7 1 

llif-liiT   fiiiii'tiiiiis   (if   (fri'linini,    siiip 

II    lull  iir  liviliiiu'ii   inn,    ItiS 

II  inn    ruiii-fiitratioM.    JJ 

after    liciiinnlia;;!',    I  IL' 
ratalvtic  |Mt\M'r  itf.  L'.'! 
tlrtrrtiiiiiatiiin    nf,   IW 
(if   iiiTcstiiial    ntiitt'iits,   .'tii.'i 
law  nf  mass  aitimi  aii<l,  I'li 
iiirtliiHJ  (if  expressing;,  'J7 
nietlHi'l    nf    ine.isni'enieht : 
electric     nietlid.l,    'Jll 
iii>lii-at(ir    inethiicl,   ;;j 
staihlard    solntiens    ftir,    .'!  1 
II  i(in    (-(niceiitratiiin    in    Meo'l  : 
elTect    en    ilissdciatiini    i  iirte,   :!"<ii. 
elTect   (in    respiraterv   center,  ;'..l.'i 
llippnrie  jicid,  ."i.'iii,   ci.;ii.   Tin 
llirihlin.    Inn 
llistainiiie.    ■\'.>7,    ."iij 
llistiiline,    C'lil.    i;i>:i 
Hdincixentisic  aciil,   'iiij.   "lill 
lldrilein,   ."i7H 
lldrindiies,  .■;,  7l'!* 

in  cdiitriil   of  circiilatidii,  LMfi 
res]>ir;itdrv,  i'.  11* 
llducll    tliei'.rv     .lildd.l    cidttiii-., 
Ilun«;er,    171 
lluii;;er    cdntr;ict  inns : 

alcdlnitie     lievera;;es    .-tntl.     t7s 
(•(introl    (if,    17ii 
(luring    starvatiiiii.     171 
in   esdpliaj^iis,    171 
inliiliitidii   (if,    177 
in   stdinncli,    171 
nerve   centers   ;inil.    17'.' 
reindte   etl'ects    ef,     171 
rlivtliniic,    171 
spl.'ineliiiic    nerve   ainl,    177 
va>:ns  nerve  and,   177 
lliirtlile    ni.'iniiineter,    iLNi,    1  li> 
llydrdceplialns,    Jilt,    ■.'.'i:! 
Hvdrdclildric  acid,   anidinit    of.    1*^ 


llypotlivrdidisni,  7.'i."i 
Ilvpiitonic  siiliition,  i! 
ll.vpdxantliine,   ii:'.."i 


and    emptying    df    stdniacli, 

fnnctidns  'df,  4Si; 

source   of.    IS.'t 
Hydro;jen  ion    i  st  i    H   ion) 
Hyperacidity,  -tlil 
llypercsfhesia,  H'A] 
Hyper^Iycetnia,      in      pancreatic 

'fiSII 

postprandial,   li."'.' 

splanchnic.   i!7:) 
llyperpnea,   :ll!t,   .'l-'!> 
llypertliyriiidisni,  7"iii 
Hypertonic    sdliitinn,    i'< 
Hypertdiiicity,   ti^! 
Hyp(i(;asfric    nerves,    7!I7 


•i  111  I 

Ignition    juice,     l;iH 

lleocdiic    iniiscles,    SSL' 

lled.dlie    spllillcter,    li:7,     Hill 

Iniliiliitiiin,    ill' 

Iniida/dle  :iiii|  (,'roulli,   ."isil,  (liil,  (ii'll 

Iniidn/ole   rin;;,  liL'i! 

Iinida/.olyletliylaniiiie,   :;:i7,    l-H,   ."lej 

liiiiiiedi.'ite   iiKJiiction,    s:j;; 

liiipiilses,  initiire  df,  s:;ii 

Index  lest,  S7II 

liidican.    i;.'!!' 

Iiidiratdr    nietliod.   li-t    ef    iiidicatins,   :'.:! 

Illddie.    ."iii|,    lint,    li.'iL' 

Inddxyl    sulphate   nf    pdtassiuin,    liiij 
,    ;:s!t  Induction,    ininiedt;ite,    sj:: 

successive,  sj  1 

liihiliili 'eciprdcal.   HI  1 

Inhiliitiiry  elTects  cf  ;iutniidMiic  nerves,  MS  t 

Innervation,    reciprocal,    s|  I 

lnor;;;inic   constituents   of   urine,   .'iIll 

liior;.'aiiic    salts   nnd    ;;ro\vtli,    "isti 

Inosiiie,  li.';!i 

Inosiiiic   aciti,   ii;i7 

Inspiration.   iie;,'.'itive    pressure   diirinj;,   :iii"> 

Integration   of   allied    rellexes.   si'i; 
tiKi  Integration    df   nervdiis   system,    suit 

Inten-dst.-il   muscles.  :;l|i 

Internal   :iii,'il   sphincter   musiles.   HSi" 

Internal    vesii;il    sphincter,    ssi' 

Internal   respir.atidn,  •"7S 

I  litest  iii:il    li.acleria,   'i'u 

Intestinal    .iuice,    conlrdl    of,    112 

Intestinal  olistriiction,    I7ii.   'I'lt 

Intestinal    secreticins.    Ill 

Inlestine: 
ntisorption   fiinn,    I.'! 
I         anastomosis  of,    I7ii 
I         liacterial   di;;estion    in.    111!' 

digestion    in.    l.Hlt 
I  law  (if,  4iiii 

I  nidvements  nf: 

larf;e,    pis 
Idi)  ;  (dinical   conditions  etTectin^,  470 

j  small,  4ii;i 

■  nature  of,  4iiii 

nervous    cdiitrdi    (if,    4()7 

Intra. ardiac    pressure   curves,    14(5,    l.'il 

Intracranial  jiresswre,  :;.'i.'! 
dialietis,         Iiitraf,'astric  pressure,   4.11 

Intrapleural   pressure.   .'104 

Intraimlmonic   pressure.   l.'!i!* 

Intra    vitan     aiiticda;;ul;ints,    Imi 

Intravascular  clnttint;.    Iii7 

llllllill.   lilil 

Invertase,  81,  4!IJ.  li'i" 

Invertelirates,    senmeiited.    7S:| 

Inxcrtini;  en/ymesj  li.'i" 


m": '.  (I 


.Pll 


i[ 


I'll 


lliviiluiltiliy    tilpiTi,   culil^r   I'l,    "'"!• 
lo.liiii'   valiio  nf   lilts,  liMS 

llllll/MtiMII,     hi 

li'i'MiiiMtioii   in  iuM\(tu?4  svNti'in.   SJtt 
IncMT-iiliility    ill    icll.'\i'<,    sin 
Isn.l.-.trir    pi'iiiit.   111 
I^i.lriiriiii'.    (ilU 

U.MIKlltciS.',    T|i 

N"iiiitiic-    luriip^l.    1  I'.i 
NiitMiiii-    snliitiiin,   I' 


.1.1.  kxihiiiii    <].ilc|i-^y.    sill 
■  liiunl.'ir  |MiKi'  ti.'hiii;;.  -7  I 
.Illirr,    yilstri'-.    l.M 

iiitrstili.'ll.    t  11-' 

]'.'iiH*rt':iti''.    1 11 

K 

K,  nil    :ni.|     I'l.'i.  k.     .niiiiii.tin;;    lissiiK' 

I1..M1I.    Is,-, 
K.'iil.  I. nil. II.    ..(•.   ls."i 
Kit..iii.-   :i.i.l.    7iis 
K,.t,.^i^.  i;s:; 

Ki.itl.'V.    i.\\;;fll    I  i-iiuil  illi.llt>    I'f,    il'.Kt 

1,111.'. v;il  '..I'.  i;-i 
>lril.-liir.'   .il'.   .'"7 
K]„v  j.'ik.   Mil.  SI.-,,   sjs 
i.iiit',.r,-.-iii,'iit    ,,f,   sj:i 

I. 

I.:i,l:in.uiiiin.  -",77 

l..'ii-t;iiii.    ill!i 

I..-i.-t:i~,',    l;il.   i;.'.7 

l.:i.ti.-  M.i'l.  •■;:'7.  ilii.;.  il7r..  ciii.-,,  7"S 

,n'i,t    1,11    n -I'ii.'iti.r.v    .•..htci,    .'.711 

ill    iti,,iiiit:nii    .^i,-kin"<s,    i',*.L' 

l,i,„lii.-..l  i.>   .■\,r,is,..  :;c.7.   11:; 

I.:i.tiiii.  111!' 
l.:iM;.ni;i;;f,    sil.. 
l.ilt.lit    l..li,,.|.    S"!' 
I.;i»-  ,,r  i;.l~.'>.  :i.;ii 

..!'   lii.'i'S  ;n*tii,ii,   :j.'l 

:i|i|.lii-.|    iiiciisur.iii.iit    .,|'    II  ii.ii    .-..Il 
tnitiiiii.    L'll 
l.i'.'i.l    |.iiis(iii!n;r.  il.'.ii 
I itliiii.    ils'.i 

I'^tiiniiti. r.    il'.'7 

ill  l.ilc,   liis 

ill  I.I. .11.1,  r.'.iii,  ii!<;i 

I Il   rNtKli't.    liiil 

I..  L'liiii.'liii.  -"7^ 
I,.  L;iiiiiiii.  "7^ 

I,,  -H.lis     ..r     n.ri.,u.     -%-l,ln.     s:;.', 

I.,  iniiic.  ilHl.  ililil 
l..ll..,.-.\t.>.    Mil 

.-.■ii^iti/iiii;  .,!'.  7'i 

traiiKitt.i'iiil,  ii7 
I..  ii,-.H-ytli.iiii:i.  Ills 

I...Mll..sr.,    Il.'.ll 

l.i'W   nil. I    ;,.,uii1i,,'   until..,!,    II 


IM>I  X 


l.iiiiiilii.i,   lii'iirtlirat    nf,    )'- 

I-i|,asc,  •_'.'>,  illi,    li»l,   CiN" 

l.ij.i'niia,   •!!•!• 

l.i|.ni.U  nf   l.lnn.l,  t'l'iH 

l.i^s'iiiiT  trai'f,    S.'ll 

List   nf  in.liratnrM,  '■'•'■'' 

l.itti'ii'.s    illapliraKiii    pliiiniiiiinnii,    '.','^1 

Livnr: 

I'irrulatinii    tlir(iii;;li,    L''.."> 

ilisnase   nf,   liJI' 

yl.vi'n^oii  ill,  fil'- 

iiint.'ihntisni  nf  f:its  in,   7i'l 

I'l  rfiisiiiii  (if,  lilH 

r.iiinvnl  nf.  111" 

1111:1    fni'inatiiin    in,   1117 
I..,.;il    irritants,   »i;t 
l,,„;iliz;(ti.>ii,   .•.■rnlirai.   Hi:; 
l.n.'kn    snilltinll,     HW 
I.iivna  ri'lli'X,  L'll 
l.nii;;s,  circiilatinn    tlir..iii.'li.   L'.".:'. 

Ml". I.'  nf  cxcansiiiii   nf,  .IJ.-. 
l.\  inpli : 

al.^nr|.tii>ti     int...     1.", 

rl.'i-trii'    .•ninlintivit.v,    li! 

lilt  nil  inn    ill,    lis 

fnrniatinii  ;iinl   .-ii.'iiint  i.ni.    Il", 

fi.rinatinii   nf,    l"i 
\.\  iiipli   s|i;ni's,   1 1--| 
l..\nii,|i:i;;.,;;in's,   1  111 
I..Mii|.liati.-s,   ll.'i 
l..vlii|.|nMytns,    IMl 
l..vn|i|inli.'  c. .11,, ills,  ilii 
I..vsiin.,  .".'.IL',  till.'. 
I,\Hiii(>  an, I   ^r.,\\tli.  .".7il 


M 

Ma.uhn  a.Mist  ii;n,  S7:i 
Maiiitniiaii.c,    dints    f,,r.    •"1711 
M.'ilincpriTs,    li; 
.Maltasn,   4!l|,   l>."7 
Mallns,,,    101 
Main.in.'ti'r: 

I.I I  uas   .lilTcrniiti.il,   .Is: 

lliiitlil.',    IL'I,    1  ir. 

iMcr.-iir.v,    1  L'.'. 

n|,ti.ali    IK! 

snriii;;,  IL'H 

valv.'.l  iii.Tiur.v,   1."il; 
Mark  time   rnllcx,    HiiC, 
Mass   aitinii,    L'^l 
Mass   n.'tinii    ami    H  Inn 
M.'iss  ninvi'iiii'iits  i.f  111.. 
Masticatinii,    III 
Mn.'liaiiiis  nf  ii'S|iirali,ni,  Jim 
Mniliiila,   sn.tinii  almv.',   s.'lli 
M.'ca.ar.vn.  ytns,  l";i 
M.'ltiiii;  |...iiil,  fats,  r.H7 
Mi'iiiliraiin  synapti,-,   7!is 

M.. y,  7Si; 

M.i.iir_\-    111:111. nin't.-r,    iL'it 


.•,.ii.'niitratinii, 
I,   I'sl 


I.NDKX 


S!t7 


:ll 


Mi't.'ilHilisin: 
cali'iilatidiis,  .'il  I 

rxii^iciKins,    *il."p 

KcniTiil,   uM 

ill   starvation,   ."liii 

iicirinal,  r>70 

(if  rarlioliyiliatrs.  il.'ii; 

(if  fats,  («k<i 

(if   [irotciiis,   ."'.'.'i 

(if   purines,   ti.iT 

s| ial,  .'31 

Methyl    ({lynxal.    li(i.-| 
Mitlivl  Kr("iu)i,   "ilts 
Metliyl   purines.   ii:;."i 
Metliylatidn,    IL'T 
MetliylKlvdxal,  'ill",  I'.C.i; 
Ntett's  nietlid.l,    1x7 
Micrdpytcs,    !•) 
Mierotoniinieter.   ."::!• 
Mill  caiiaiity  (if  liiii^'~, 
.\lill<,  cldttin};  (if.    l>s 
Miniatur(>   stdiiiaili,    III:'. 
Miniin.'il   air,   .'.dd 
.\riindnuili';ir    lencii.-\  tr^-,   IMi 
Miirawitz   (1 y.   Iij I   ilntlin;;.    Id- 
Motor   arc-is.    alilatidii    .if.    HH\ 

sliiniilatii f,   HH,   s|i!.   HIS 

M(it(ir    nerves    of    segmental    dint    iiiiiscli' 

sm 

Miiiihtain   sii-kticss.   .■'.tin.  '',W 

aila|itati(iii    tii.    |i"l 

alveiilar  Cf),   in,  IttiO 

lildiiil    ciirpiisiles    in.    Mil 
MiiMiiieiits,  (if  intestine.    I'l.'! 

(if   stdiiiaili.    1.":; 
Miiiiiiilial   fdd.l   statistics,  ."HI 
Miisc.aiiiie.  ai-tiiiii   (in  lieart.  L'U'i 
Muscle,   c.-iriliac.    prd[ierties   (if.    17'i 

refractiiry   peridd,   17H 

respiration   in,   dO.'i 

staircase    pliendinciidii     Iti'eppe'.     17 
.skeletal,    177 

respiiatidii    in.   .'»!♦( 
Muscles,   .inta^iiiistic.    HIS 
Muscular   exercise,   l.'!'..   .".".1' 

circulatdiy    cliaiijies   iliirin;.'.    l|d 

effect  on  iiii'talidlisiii,  ."i."il 

effect  on  respiratimi,  ."ifirt 

H-ion  duriiij;,  41.'{ 

purines  during;,  (it7 

redistriliulidii   df  lilddd   duriii;;.    Il'i 

respiratory   clian;;es  diirinjj,  410 

temperature   df   lildd.l    diiriii;;.    ll'i 
Mutual    precipitatidii    of   cullniils,   ."li 
Myenteric  reflex.  7I'(! 
Mydjreiiic   liypiitliesis   i.l'   luMrlliiat,    171 
Myxedema,  7."i.") 
eiii'rc.v  output    in,  ML' 


N.nvi.lics  and  lilimd   fat,  CHS 
Xecnisis  (if  liver,  020 


.\('t:atiM'   |>ii'-~iiii'    ill   \iiitricli',    l.")j 
N'eplicldincter,  ti;i7 
.N'eplirectomy.  til'l 
.\(  pliritis,  li."i(i 

acidosis   in,  lis.', 

urea   retentidii   in.  ."ijs 
\er\  ('s : 

(if  skill.  7l'ii 

net  work,    I.   L'O 

ri'L'i'iieralidii   of,   liO 

sc;,'iiieiital    disti  iliiilidii    df.    s:i7 

~|ii'ci(ic    pcdpertics   iif,    7S'i 

\.'iMi.|il:it(ir.    707 
\i  Tve  cells,  :::',,  70!t 

N'er\i     ili^ellf  rs.     1^.:  1 

Xerviiiis  cdiitriil : 

('(   ;;asfric   secretlnii.    l;il 
df    ilcdciilic   s|.|iiiictrr,    Ills 
df    intestinal    Klaii.i-,    I  II' 
df   intestinal    imiM'tiicMl^.    \*'>7 

(if    pMllcI'e.'ls.     tJ7 

df  sali\ary   yiainN.    IL'-'I 

(if    stdlll.-lcji     nidVi-lliellt»i.     t'lS 

N'crviiiis  ilial.itrs.  r,7L' 

ill    IIIMll.    ti7  I 

\cr\iiiis   sy.^tein  : 
aiitdiioniic.    S77 
liiilliar   (iliers.   ssj 
fiiiictiiiiis  (if.  SSI 
sacral    tiliers.   SSl' 
tliiiracicdliiniliar   lilu  r«.    ssn 
elTect   (if  section   at    \;iridii>   Icm  K  ,.( : 
aiiteriiir    rmil.   W.    s.',." 
Just    lieliiihl   niedulla.   si'.O 
just    lieliind   pust.   ciirp.   (|iKi.|,.   spi 
just    ill   fnuit   df  ant.   corp.   i|ii.'iil.,   s|i 
|"isteriiir  rmifs.  s;i(l 
spinal   cor.l.    s:;;i 
(■viiliitidii  (if.  7ls 

illllllellci>   (111    e\.-l'eti.iii    (if    liiilic,    ."ij'.i 
iiite;.'ratidii   df.   7s(l.   soil 
N't  work.   inr\c.   7'.ni 
neurdfiliiils.    sdn 
neiirdpile.   7s|,   7!i7 
Veiiid^'enic   liyp(itlie~i-,   i.f   heart.   17l' 

Nelll s.    7s  1 

assiiciatiiin.   7h:1.   7s."1 
iiiternieiji.ati'.   sit;.| 
iiiteriiuncial,   Sou' 
\ciitr;ility.   recrulatimi   df,    :iii 
Xiciitiiie.    -J.!;! 

a.tidii  dii   valgus,   L'L'ii 

\is<l    li.i.lics    Sdll 

Nilniy.  ii: 

e\creli (.     |.|r I;, I     I  i^r,    :,i;ii 

ill   stai\:itiiiii.   -'iii'i 

iiiideleiniiiied.    in  inc.    (i|:: 
Nitrojicii   li.'ilaiici.,   ."i7d 
Xitrdp'tidiis   ('onstitiients    i*{    uiiiie.    '<:!'' 
N'itrojjeiidiis   e()iiililiriiim.   ."i71 
N'iti  ii^eiidus      nictalidlites,      in      ^taivatidii 

.'iliH 


SUM 


INDKX 


i  t   "f   :  'I 


■k 


Nciciccptivc.    7!P.".,   HiPt 

iiii|Milsrs,   h:(l' 

ri'tlcx,    ML'.'i 
N'liiihl   vital,  '1-7 
N'oiirlri'trolyti's.     \i\ 
Niiiitliri'sliiiM   siili^Iami's,  ."ill.' 
Ndiiii  il    ;ii'ii|,    -- 
Niiirnulilasls,  !'.i 
Nllrll'JlSl',    liiiH 
Nii.lrir   arl.l.   iW' ,   iNlt 
Nihlriii    fcniiriil-i.   I'll 
Nurlrins,   ii;'.7 
\ll,'lr,.si.|,',    t\:\H 

Niirlc.tiilr,  i:::>4 

Nvsta^iiiiis,  h71.  *^7'i 

() 

(HH>it\,   Haiitiii^'  iiiii'   fur,   "i7I 
Oleic-    iiii.l,    MlH 
(tl.iii.   >*Hs 
npsdiiitis,  7i* 

(»i<;aii<.    liis<   "f    wi'i^'lit    ilurin;;   statvatiiiii, 
."ids 
|,frt"ii>iiiii   'it'.   •'Is 
Drnilliiiii-,   I'll'!.   <'':ll 
(liiiilliiiri''   aiiil.  'i^'.l 

iirtiiii|iii''a.  :;i:!,  :'.is 

Osiillat'.ry    ni''th"il   "I'   lil""il    |ii''>snf,    Kl" 
()viiinin''t''r.  .'i,   L'.'.ii 

()<lll'isi<,     1 

llsrii'itii'   pri'^s  111 

ami    fiTnia   .  if    lyiii|.|i,    I:; 

:iiii|    lii'iiiiilvsis,    7 
jiiiil   iilasiTi'il,\si<.   s 

iia'aMiii'iniiit    ii.v    i|.'|iri-si'iii    nf    (iff/ 
iui:   piiiiit.    1 1 

in     jilivsidlii^'ir     I ■Iiaiii-'iii-',     l.'i 

ill   piii'liiiti'iii   "f   luiiii-   liy   kiiliicy-'.   11 
iif  traii>fu>at'>.   1  II 
( )\altiiiiitiii.    :i^    fi'iiil.    "'77 
(hiivit.-llin.   a-   fiiii.i.      77 
0\i.lasi'M,    SL' 

Dxi.lati'iii     'if    I'li'ii'l.    :'i'»7 
Oxyl'iitvri.-   aii.l.  I'lii,  Us:;.   7'i!i 
<  txyycii : 

'•iifl11ii''iit   "f  "xi'Iati'iri,  .'!!':! 
'IctiTiiiiiiati'iti   i.f,  ."itii; 
'■stiinatii'ii  in   lil'inil,  li'.Mi 
i<'i|iiiniiii'nt<    I'f   tissui'.*,   :'.;'■". 
tiTsi'in    ill   alviMilai    air,      .i-lii,   .'Itt 
tciisidii  ill  aiti'iial  IiI'mhI,  :'i.'17 
tr.'iiiNjiintatinii    by   M'to'l.   .'t71* 
vcliinic   I'di-i'iiray^  in  lil"0'l,  :Wii 
()xy;.'iii    iiiMiifti'ii'Tii'y,   .in'l    ]ioriiiilii'   liri'atli 
inn.   •'"■' 
ifTi^i't    of,   on    ri'S|iirati"ii,   :i."ii,   !!."!* 
(Ixypii  supply  "f  'iiart.  Kit 
()\ypi"t''i.-    .■i.-iil,    <i:J!t 

!■ 

I'airlii'iniaii   lin.ly.   Jl'.l 
I'ain: 

sciisati'in  "f.   s:'.i.' 


Piiin.  Hcns.'iti'in  of- font  M 
transmission   in  lonl,   H.'tO 

SIMIMP,     7i*.'l 

Talataliility,  ."lit;! 
I'almitic  a''i'l,  tiHii,  7ii7 
I'aii'ri'as; 

liiirniiini'  i'"ntr'il  "f,    IL'" 

liistoli'^ic  '■liaii;;''s  of,    !-',» 

(ixyyi'ii    n''|iiiri'iiii'nts,  :'.'.••! 

n'Tvius  ''iiiti'il  "f,    t'.'7 

siij^ar  nit't:il"ilism  ;in'i,  tt7H 
I'an'Tcati''    'lial"'t.-s,   t:7s 

1'; ri-ati'-    'litf''stiiiii.    Is'.i 

I'aiirn'ati''    jiii'M'.    I II 

ami   fat   .iiui'stion.  ilmi 

sf'-ri'tion  "f,    iL'o,   l-.'ti 
I'aii'ri'atin,     ll'H 
rarasympatli''ti'-   systnii,   ssl' 
l';irii\ysmal    ta'-liyai'li.-i,    -'''!',    -7H 
I'arlial    'liss,ii-i:iti"ii,   J71 
I'artial    pn'ssurr   of   y;iscs,    :',:',!'< 
ratliH;iys,   si'iisory,   in   spinal   'Mini,   Sllii 
I'olvi'-   (lanijli'inii-   Ki'mi'.    ''"^ 
IN'ntosc,    titi4 
I'l'lisin.   a'-ti'in    of,    is.'i 

]irnilufts   of,    isti 
I't'psino^i'n,   4S.'i 
i'.'ptiil'H,  tiol 
l'''ptiim',    III".    IHIi 

I'lT'l-ptilill,    Hill 

rirfiision,   of   kiilmy,   Hil 

of   livor.   Ills 
Perfusion    Ihiiil,   of   li'-art,    I'i"' 
Perfusion   of   heart,   UU 
Perio'lii-   lire.Mtliiiic,  (•aii^i's  of,  :i7i; 

typ.'s    of,    :171 
PiMi|i|ieial    ri'sisfaiii'',    PU,    --I' 
PeristaKis: 

in   esiipli.'i^iis,    t  Iti 

in    lar^'e    intestini',    I'iS 

ill   small    inti'stini'.    111"' 

in   stomaih,    I"'.'!,    t."'i' 
Pirist.'iltie    riisli,    Itlll,    1711 
Perlstaltii'   wave.    111."' 

PiTiiieioiis   ani'inia,   I'liiTny    output    in,   ."'4-' 
Peroxiilases,   s:^ 
I'M,  -.'7 

Plia^oeytes,   !*7 
Plienai'Ptnri''  aeiil,  710 
Plii'iii.;,   ."Ill 

olai'etie   ai'iil,   -"H- 

plitlialiMn,  4S-.>,  .",:;.-| 
>lae'^i'-  aeiil,  liiU,  71ii 
Plieiiylalanino,   1104 
Plienyl  iiroiip,  T'lM 
Plilorlii/.in,    titl4,    tili'i 
Pliospliatos,  oxeretion   of,    17 
Plio.spliatc  solutions   for    1!  ion,  M 
Pliospliatcs  of   urine,  .'.l- 
Pliospholipins,  tlM't 

in  I'ilo,  4!tH 
Plirenii'    leiitcr,   -i-H 
Plivsi'oelienii'iil    Imsis,    1 


INDKX 


899 


f 


VhynUAtni'ii-    priiicsMos    ili'|it'iiiliii|;    on    ail- 

snr|iliiiii,   ti:t 
Pinnipiifs,  aliHor|ifiipii   of,   117 
I'Mocarpiiu',  in'tidii   mi   lii'art,  '2M 
riliinicitor    fihiTs,    HHil 
Titot's  tiil.rs,  :;iil 
riiisma.  !'!• 
I'lasiiKilysis,  S 
I'liitolc'ti,  'if  IiIimmI,  !i7,  liMi 
ri.tliiira,    Kti 

rii'tii.vsiii(it,'riipii.  i;it!",  '.'.MK  'JT:'.,  :tii:t 

I'li'iirisy,  .'!J4 

IMi'mim"iiI'  AiictI.mcIi  :iii<I  Mcis-iici.   HH;.  T'.'tl 

riiciiiiKitliiiriix,   :!ii." 

I'dikiliHVti's,  !M 

roivurai.ii,  -::: 

I'lilyiu'uritis,  .'iS) 

I'olyiiiiili'ar  ci'lN,  '.Mi 

I'dlypcptiilis,    IH7,  tiiil 

riily|ili(is|ilinric   a<iil,   <>'■''' 

l'..|v.ai'i'liarh!i's,    Is'.i 

l'(ilvs|pliy);iii(ii;raiii'<.    -7:1 

I'lirtal    vein,   Moiultlow    in,   '^"t 

rn^tiliiTotii'  wave,  |iiils('.  L*";: 

l'cist|iraiiilial    liypc  r^'lyii'iiiia,    lil!' 

I'l.stcciitral    riitivcilntiiMiN,   H.-(i,   H.'i  t 

rostt'rinr    ntnts,    7H7,    X'.U\ 

l'nsts|tliy^ini('   jicrifni.    l."in 

I'dstiirai   rrllixi's,    HL'ii 

Pntassiiini,   niirr«H*!irinic-al  ti'st    (•>i\  4-1 

l*ntassiiini   iims,  on  licart,  l<i7 

I'dti'iitial   aiiility  <if   iiriiic,  rt2i 

rrcct'iitral   coiivdliitions,   HC!,   H.'il 

rii'ii]iitins,   "ili.'i 

I'rciliiTdtii'    «avis    |iiilsi',   L'ii:i 

Prefrontal   rcjiiori,   h,"i  ( 

rri'iiiatnrf    heats,    1177 

rri'inortal    rise,   ."ii'iii 

l'res|iliy;;nii('   perioil,   1  !!• 

Pressor   imiiulses,  :;:'iS,  :;;!!•,   Jlii 

Pressure: 

intrapleural,    ^'.nt 

I'ffei't  of,  in  liloiiil  pressure,  lloii 

intrnpiilnninie,  *JW 

negative,  llo'i 

(tsniotic,    1<> 
Pressure  pulse,   127 
Prineiple  of  \Vill;ir.l   (iil.lis,  (!(1 
Proline,   tiOJ 

Proprioeeptive   impulses.   S(i." 
Prnprioeo|itors,   8:;J 
Proseeretin,   4'.'<> 
Proteases,  8!i 
Protein  sparers,  ."i71 
Proteinases,  Hn 
Proteins; 

as  eolloiils,  >'<■> 

baeterial  distestion  of,  ."lOl 

flieniistry  of,  ."1>7 

metabolism  of,  ."O'l,  fil.'i 
eiiil    proiluets.  tilH 

minimum   re(|uirement,   ."72,    jOl' 

of  bloo.l,  88 


Prnteins-C.oit  M 

ri'lative  value  of.   for  (;ro\vtli,  (ill 

milting  out   of,  till 
Proteose.  4HH 
Protopatliie    im|>ulses.    Mill 
Protopatliie   reieptors.   7!io 
Protothroniliin.    lo:i,    imi.    111 
Psyellopatlloloyy,    8(12 
Ptonuiines,   ."iii2,   tl2!l 
Pt.valin.  4ii|.  ti.'iii 
Pulminiary  einiilation.  -''■'■ 
Pulnionarv   ventilation,  .'l.'io 
Pulses,   ll'm 

abnormal.  27*1 

.•ilti-rnans,    I  si 

io^eniirnis,   Isl 

enntour   of   \\a\'e,   2"0 

len;;tll    n(    v\:.ve.    \W 

palpabl,..  2o| 

pressure.   12. 

|iulse    ilirves,    202 

pulse   Haves.    1S!I,   200.   2o.l 

rate  mJ'  transmissiini,   ll'H 

veloeity,   2110 

\enous,  ei-ntral,   2  ■■'•,  271 

venous,   peripliei-al,   2o.'i 
Purkin.je   lii.ers,    |sl 
Purine    boijies     i  .v<  <      I'inines  i 
Purini's: 

clieniistry  of.  .'.2!i.  Hi:'.,   il:i  t 

enilo»renons,   till,   *H:i 

exojxenous,  (ill 

tnetabolistu  of.  tl:i7 

in  starvation,  -'til* 

synthesis  of,   llltl 
I'utri'fai-lioh,  intestinal,  ■'.ol.  .",:iii 
Putresiine,   ti2!» 
Pylorie  canal.  4."2 
Pylorie    splliuetlT.    eontrol    nf.    I.'ili 
Pylorii'  vestibule.   4".l 
Pyranii'lal  ei'll   lamina,  H."  I 
Pyrimiiline  bases,  tillli,  li:!7 
P'vruvi.-   .-oi.l.   lion,   7ns 


Kami  eommunieantes,  2.'i:i 

Rayuaml 's  ilisease,  blooilllow   in,  2'iS 

Keaetion  ileviatlon,  871 

Reaetion  of  urine,  .T24 

Reaetiiuis    ilepeuflin^    on    a*lsorption.    fitl 

Keaetions  of  biuly   lluiils.  :i." 

Reeeptors,  782,  7>«>* 

<listance,   78." 

epieritic,   71»(i 

extern.al.  78S.   s22 

internal,  788 

of  skin,   7!ln 

projieient,   78.",   7S8 

proprio,  822 

priitopatliie,  7f'o 

temperature,  7!U 

tcuieli     7O0 


!MIII 


iM>i:x 


III'    1 


li.'.  i|.rM.';il    iiilliliiti HI  I 

rictiiin    lit'   itrM'liiiirii'   <iii,   HI!' 
Itri'i|iriirlil     iiinrrviitiiiii     nf     lilmiil     vissi'N. 

•J  1 1 .  H I  I 
l!i'i|    liliiml    I'lirplisrlr-;,    ii!ii;ili    iif,   !'i; 
K.'iliictiiiii  of  hliMiil,  :'.h7 
Kil'irrcil   piiiii,  HH.I 
|{illo\.  11111. litiiiMi'il.    i:il 

unritn<litii)!)t'il,    \l'<] 
KilliA  :irr,  7st 

iiftiT  ifTivt,  Hln 

;;r:iiliri;.'   nl'   iiiti'ii»il.v,   Mi;t 

iMrvrr>iltiIit_v    i.r    iiHi'lii.timi.    '*1" 

l;clrlit    |iiTii.ii.  Mill 

it\vj;('ii  lii'j'riMititiri.  Hi:; 

|irii|irrtiiv  111',  t:{,  :;!i,  in 

irt'nii'tiiry   luirioil.   H)  1 

^iiiiirii.'itittit.   H)(t 

l.'iMlrx   luiti n'<i-l:ni.i'   i.f.   Hi.", 

UilliAis: 

allii-il.    ^iiniilt;iiiriii|.-    inti'ijt;iliuii    nf,    hlM 

;nit.'i),'"iii-lii'.  h:;! 

:i\iiii,   "!'" 

li.-iliiiiski,    Hu; 

innilitiiilli'il,    H."ili 

rrrniiiitrrii',    S-lfi 

rni^snl    rxtrllsiuli,    Hii| 

rxtrtisiir   tliru-it.   hii.'i 

t':iti'_'iii'   lit",   Sl'". 

■l.'\ii.M.  Hill,   Hjl 

iiiti'i;riitiiiM   ■■!'  .■illii-.l.   »:JI 

intrnii'tinii   littwrt'ii.   H:_'i 

in:ii!i:itiuii   nl',    h-jiJ 

mark  'inic,    Hiiii 

iii,V'  iitrrir,    71"! 

!,.'itiiri'  iif,   S'J." 

iiii.i.i|ili\i',   Hjr, 

liiistural,    HL'i'i 

iini-iiii.litiiin.'il,    i:;!.  H.'iil 
Uil'r.-iitivi'   iiiili'X,  liloiiil.  HH 
l{.  fiailiir.v    iicriii.l,    HH 
Ifil'ijii'triinii'trii'    iiirtlioils,    HS 
[{.■;,'iiiiMatiiiii   lit"  (■r.vtlir(ir,vti'«.   li:i 

|{r;.'iilati t'  iiiMitralit,v,  ;:il 

|{i-;;iii';iitatiiiii  of  ;;astiii'   riiiit.Mit-.    I  l;i 
Ifiiiliait  Mi'i-sl   value  nf  fats,  liHH 
Ki'infiiri'iiiiciit   iif   kiH'c  .jr'k,   HJll 
Kiua!  ilial.cti'S,  filil 
lii'M.al   fiMutidii.   tlifiiiii's   iif.   ."ill 
Kinnin.    »HS 

Ki'^iTvr   alkaliiiil.v,   iniasuri'iiiriits   iif,    imli 
lilt   mi'tliuils,    I'J,    U! 
nil  a-iiri'iiirnt   of.  Mtriitiiiii   iin'tjioils,    II 
Hi'>i.|aal  :iir.  .".(in.  ".1 ! 
l;i~liiiatiiiii: 

aii.liiMiliial.   :;ii7 

lii,vuic.l   tlir   1iiiij;>,   ';7"> 

iliiriii;:   iiiiisi'ular   rxrnisr.    Ipi 

in   riini|iii-isril  ail',    ln- 

in    rarrlitil  air.   ".'.I'.i 

nii-i'li.'inirs  of, 

inoM'nifiiN  nf  .-lit:!;:!!!   in.  .".21 

nLiiviMiii'iiN   I.f    I  ,1.^   ill,   :'.l!t 


iniiM-nl.'ir   rxriri.<t'. 


ilit,   .■i.'.ii 


HI 


■<|.ii'jitiiin  laliiriiiii'tir,   'I'M 

^piriitorv  iriitir,  ilL'7 
iifTcri'iit   iiii|.nlsi.H  to,  :i:'.l,  IliiU 
niitoniaticitv   of,   '.'rM 
liornioiii'  I'l.iilriil   of,   :!.!.',  .".Ill 
rctlcx    control    of.    ",:il 
sonsitivit.v  to  alvrnlar  ("<)  ,  :i."i7 
stiiniil.'ition   li,v  CO.  ^l.'ii; 
xiilwiiliar.v,    ",-H 

s|iir;itorv   ili;ni;;<'s    in 
(ill 

'-|iiiiitory  iMlianyo: 

; r.lin;;   to    lpiii|\ 

ami  lioily  tiiii|.i'ratnri',  .".."il 

I'liiiiral    ini'tlioil    fur    ili'trriiiiniiit;,    '■ 

ill  ilialutcM,  t'i7H 

ami    mn«iMil;ir   I'Mriiso,  .'.."il 

.•mil    ti'tn|irratiir('   of   mvironnn'iit. 

in   tissiii'.*,   :i!i:t,  .",117 

's|iir.ator,v   liiirinoni.,   iiatnri'  of,   "111 

•spir:itiir,v   moM'inrnts,  .'11.'! 

■spiraloiv    pa'siiyos,    prc^stiri'    of    air 
L'lili 

■^ptf.ator,v    i|iii.tii*iit,   ."»l."i 
in  ilialiitcs,  i'.7H 
inllucmc  of  .lift  on,  .1 17 
intliK-ni'o   of   nii-t;)lMilism   on,   .'ill) 
inlliii'iU'o  of  iniKiiil.'ir  .'utivil.v   on. 

>pirator,v  trai'in^s.  .'{n.". 

.*pir;itor.v   valvi-),   lVari<'>.   ."iM 

tiinlati'il  eiytlirolilast^.  li.", 

Ai'fsil.lr  action    of   cn/.nnis.   :.'."i 

lis,    inovcinonts   of.   ill.'* 

miisinlaturc  of,  oli> 

iiniliil.'itor,v   niovcineiits   of.    ".17 

t;lit    lateral   coiinoctioii.   Inart.    Ih.'i 

yiilit.v,  (Icc.Trbratc.  huh 

ilamlic  lissnrc,   h."i."i 

lots,  787 

anterior,  7h7.  h:i.", 

posterior,  7h7.   hih; 
vtliniic    sc''inent;itii 


i.-,l 


111, 


.' » 7  " 


Hi  I 


S:o-ial   niillluu.   »■-' 

Saliivlates,  liln,  ii.-,7 

Saline   injection,   efTeit   on    lilooil    presMire, 

i:i!t 

Saliv.a,    control    of    secretion,    nervous,     IL'". 
p^vdiic.    I"l.  H.'iii 
iionnal   secretion,    t.'ll 
Salt,    ilietetic    value,    .'.Hil 
Salt.'il   lilooil,   loll 
Saltiii;;   of   protein-.,   tin 
Sapi.nitif.'ition.  i^h7 
Sarcosiiie,    r,L':i 
Saturation   limits.   li.'iL'.   i'..". I 
Scratch   ri'tlex,   Hii.",,   .H-.'l 
Scurv,v,  ."iH."i 

Sea   anenioiip,   nervous   svsteni   of,   7H.1 
Scion. 1   wiml,    11. "i 
Siiretorv   tili.'rs.   varieties   of,    IL'I 


iVi\ 


l\l>l  A 


'H)| 


III, 


S.MTllM.      IJ.". 

ilii'iiiical  iiatiii'.'  Ill',    ll'ii 
iiio'lianUiii  iif  action  of,  4Jii 
Siirctiiiii    (.>»(•  uihliT   vurinu<<   (jlaml 

KiMi'ral    I'dnsiilirations,   -llS 
Si'uhu'iilal  ilidtriliiiticiii  of  iicrvri,  H: 
Siyiiu'iitntioii    inoM'Mii'iiti,    lt'i:i 
Sr;;Mtflitril     III vcrtrliral t"J.    ticrviiii-*    s 

Soiiiii-iri'iilar    tanaN,    h7;'. 

i'Vf   iiiovi-iiu'iit^   an<l.    "*"' 
rrmoval  of,  H7  I 
Si'iiiilimar  valvi'S,  I'lii.   1,",", 
Si'iiii|iiTiiiialil('    ini'iiiliiaiif.     I 
Sense,    t<'iii|p('rafiiii',    Tl'l 

toiiili,    7!i:i 

pain,  T!>." 
Sciiiory  ci'htiTs,  S.'ii.  s.', | 
Seriiii',"  (in:', 
Siinin   nlliiiniiii,   s7 
SiTiiin  uloliiilin,  H7 
St'v,   ifTnt    on    iri'atinini'   oxi'iction, 

cfTi'it   on   iiiir^jy  output,  "II 
Sli.iiri    ffiiliin;,    •(:!.') 

siieii  sihhk,  ;;h7 

Slio.k,    l.'S7 

.■inestlii'tii',  2SS 

lilooil  pri'ssuri'  in,  Llim 

cxpi'rinicnt.Tl    invcstiuatioiis.    C)>*!' 

1,'ravity,    2S7 

licnwirrliaiiic,   2SS 

nervons,    2S9 

recovery   from,   S(i." 

seconilarv  sviniitoins  of.  '.'!•" 

spinal,   2S8,'  Ho.! 

siiriiieal,   2S!) 

treatment   of,   20." 

vasomotor  eontrol   In.   L'!>0 
Sinoanrienlar   no.le,    IS.",   'JiiH 
Sinus  arrliytlimia.   L'*>*i.   1.'77 
Slims  loailvi  .■inlia,  L'tiil.  '.'77 
Skatole,   .-,01.   c,:;^ 

in   urine,  .".".I 
Skeletal  miisile,  ii-|iiration  in.  .'lOI 
Skin,   reeojitors  of.   7!>o 
Soap,   GSfi 
So,llum  ions,   liJii 
Solution  of  aases.  :;;;il 
Siiliitions: 

was  Laws  ami.  .'1 

i;rani   inole.nlar.   .1.   'JL' 

liypertmiie,   liyp>itoni<'.   .'iii'i    ismoni. 

nature   ot'.   :i 
Siirensi'ii     metlioil     for     est  imatiiii,' 

y:roups.  .-.im 
Soiiiiils,   c-iniiae,    1.".7 

riHorilin^  of,   l.'S 
Speiilie   concluetivity.    17 
Speilfie  liynanili'  aition.  ."i:',"^ 
SpecKie   j;iavity   of   urine.   .'iL'L' 
Spliiii);omyi'lin,   (I8!l 
Sp!iy;xmi''    perioi],   27.'I 
S|i|iyi;llon;rapll,    Mihlt; 's.    'Jitl 


'spinal   aiiiioal.    ^"  | 

S|iinal  eoliimn,  7^! 

S|i|nal    eonl: 
»i  alilation   of,   H.ili 

in    latioratory   aiiimaU.   ho:; 
I"  III  man,  8mi 

liemiseilioll    of,    s:;l 

.^sleoi,  siiisiiry    patlm.avs   in,   .s.tii 

s essivi'  .leyriH'r.'ition   ill.   ">i:; 

spinal    pathways,  alTeri'ut,   "»:;■! 

>piiial  slioek,  Hii:;,  ho7 

spirometer.    .""I'i 

>-plaiii  liiiif    I'ireiilation    in    sliock.    L'112,    21' I 

>pl.iiieliiii,'    nerve,  2;i:!,   r>72 

>poii;;i's,    nervous   Mystem    of,    7^2 

St;il;i^mometer,   li."i 

Slamiaril    nf    iieutr.-ility,    2ii 

stamlanl    solutions,   jiiep.'ir.at  ion    of.   .'■.I 

staiiiitiis*  li^jatiire.   I'll 

starvation,  ."till 

aiiilovis  iliiriiii;,    ''l<'.> 
r.L'l  laii-e   of   ileath.   .'.711 

cITeit    of   ireatiniiie   exeretioii.   t'lj", 

iMi'i^y  output   iliiriii^,  ."iiis 

ivi-retioti  of  nitrogen,  ."liii 

I..SS   iif   Hi'lylit,   .'ii:s 

iiitroyeiious    inetaliolisin,    ."iIH 

juiriiies   (lurin;^,   ."l»!l 

secretion   of  Kil^*rie  Juice   tliiriiii;.    I7il 

si'iisations    iliiriiij;,    I7.'> 

sulphur   iliiriiin.   ."iliit 

In-.atinent    of  liialietes.  liSl 
Statistl.al    metlio.l.    in    ,liet    coiitml,   ."H!l 
sti'aric   aciil.   i!s7 
-Stokes  .\i|;inis  s\  iMlrniu.',   IPIl 
St .-oli: 

.'irranueiiient   of   fimil    in.    1."' 

'!i«;estion    in.    1^1 

elll|ityill','    of.     I.",il 

elTlct        of       patholnoic       C litiollS       ,ill. 

Ii;m 

late    of.     (.".S 

;;.is    In.    Iil2 

miniature,    i'.V.\ 

nio\enients  of,    l.'il 

efl'i'ct   on    f 1.    I."il 

Stionia    of   re.l    cell.   PI 

stromulir.   2ii7 

stivcliniiie.  .'ictiiiii  III!   reci|,incal  iiiliil'itlon. 

sill 
Suh.-iracliMoi.l   ^p.-ice,  llil.  2IS 

Sill still  allele.   :;2I 

Sul.costal    l.oi.lcis.   :;l'1 
SulMliiral   space.    Ili; 
Siiliiiiicrons,   ."il 
Siilist   ntia-t!el;itinosa,  S;!l 
Successive  iiiiluction,  ,S2J 
Suc4'essive   re^reneratiou,  SI.'! 
Siijjar,   storage  of,  liiJi' 
Su;jar   level   in    lilooil.  I).~i7 
SM!;;ar     metalMiiisiu      t  .\r  r      t'arlMiliy.lr.'ites  t, 
l!.'2 
r.l.-ition   of  pancreas  to.  ii7S 


fH)2 


INKKX 


;  ^i. ' 


r^* 


irifi  •! 


Sul|>li:iti'8,  ithrrcal,  ii::'J 
Siil|iliatis,  iif  iiriiii',  .l-'lj 
Siilpliiir,    rxiritiiiii    i<(,    '11  I 

III    ^t:lr^llti"n,    'li!' 
Siiinni.'itioii    III    ri'lli'M'".    ""I" 
Sil|M'riiir   lar,vnui';il   ihim'.  iiitliii  iiii'  nil 

|iiriitiiiii,    X'•^ 
Sii|>|<li'iiii'iitiil   iitr,   iliiii 
Siirt:hc   iirrii,   iiikI    riii'Ufv    imtpiit,    ."it 
Sitrt';i<'i-   Ti->i^inii,   iiii-iisiiri'iiH'iit   nt'.   *il 

SiirMViil   iMiin.l,    ■hii 
Sii>pi*nsi(inM,    ."1 
Sii^|ii'iisui.|-..  .■.•Il.iiil-,  ilii 
Swnlliiwiii;,'.    ll.'i 

iiiit.r,     117 

iif  lii|iihl   t'iMiil.    IIH 

lirrviiiH   inliltipl    nf,    I  17 

•iipuiicN   pni.liiir'l    liv,    I  P.I 

X  riiy   iliiriiin.    I  'i' 
S».';it'  (,'l:iii.l«,   SMI 
Sviiip:illiflii-    ciiiilicil    !•<    Iii':irt,    •J:;7 

"!iff.T..iit,  •:■:■: 

Sviiip:itlii'tif    iiiTvi',    IJI 
sViiipatlii'tii'  sv^ti'iii,  N7'*.  ssii 
Synapsis,  7x1,'  7!'7,  s|!» 
Smiiptic    fatiyiir,   '."."i 
Syii.'iptii-    iiii'inliraiir,    7I'"« 
Syiitiiiiiii,    IHil 
Svstnli,.    ill.lcx,    iL's 
Sy»*tolir     pn-sslllr,     1J7 

iin',')<m''iiM'iit   itt".   ill   mall.   I'JS 


T,mI"'»  ili.rs-ilis   -""i 
'I'aclivi'anlia.    paii'W-iiii.-il,   'Jiil' 

Tai-tili'  iiiipiil»i-s,  s:;:'. 

traiisiiiis'^ittn   in   »-iir<l.   s;;;; 
Taiiriiii',    P.I  I 
TannHliiilii'   a.i,l.    P.i  ( 
'I'riiipt'ratiiri* : 

iiftrr  itTi'.t.   7!»L' 

rtV.-.t   on    iiirtaliulisMi.  .'"l 

■-iMisati.iii   lit'.   7I'L'.   s.;!' 

traiiNiiiissinii    ill    i-otil.    s;tj 
Ti-rnpiirary   assi»4-i;iti(ni.    ^,"17 
T.iiiloii  .jirks,  SI'S 
TiMisidii   of  CO,   ill   viiMMis  I.IimmI.  :!lL' 

nr'   nascs   ill   aUi'iilar   ,'iir,   Hi,   :!:'.!• 
Ti'taiiiis.    ill    stciinai'li,    171 
Tct.'iiiiK    tiiviii,    a. linn    nil    iiiiprni-al    iiilii 

liiti >I!l 

Tliiini'.  il.'i.'i 
Tlii'iiliroiiiiiii'.    I'l:!."! 
Tln'riMiii"<tlirsiiinirl<'r.   7!il 
Tliiiraiic  iii'cri'uliiin.  .'ill! 
Tliiicai-i.-Hluiiiliar  (nilllnw,  SHO 
Tliriinibiri,   I'lJ 
TlirotiilMPjini,  liiti 
Tliroiiiliiikinasi'S.    Imi 
TlinmilMpplastin,   UMi.   Ill 
Tliriiiiil"i>*i-i,   1117 


Tliriiiiiliu     fiiriiiatiiiii,   ll:i 
Tliyiiiir  ni  111,  <>l!» 
Tlivmiiii',  •l.!7 
Ti/lal   air,  Mii> 
ris-.Mt    nii'thiiil,  .'ill,   .'i.'iii 
Tls<<iii'   lliii'l,    I  l*i 
Ti'Kiii'    jiiii"',    117 
TisMifN : 

niiiliKi  ai'i'U  ill,  <'<'*7 

iiilliii'iiic  <it',  on  riottiiiu,   lilt 

(•xyt'i'ii    ri'i|iiiriiiiriit!<   of,   :i!>:i,   .'i!»7 

iitili/atioii  of  >;liiroHi'  liy,  liHl 
TitiaMi-  ai'iility  niol  alk.'iliiiity,  JJ 

T iii.'tiT,  :i:iH,  :isi 

ToiiiK  rliytliiii,  of  Htoiiiaili,  t71 
Toniilar   lirropliili,   lI.'iO 
Toll!  Ii,  i|i:<iTiiiiiiiatioii,   7'.'t 

lofali/atiiiii,  .';7,   7!»."i 

Nfiisc,    7I'it 
To\iiii,  tilt 

Traiiifu«ioii    of    liloo.l.    1:1.1,    i:;!t 
Trrpliiiiinu'.    -'•■^ 
Troppc,   17<* 
Trirliloilarlami.il',  tl.i.-i 
Triiiii'tlivl.iiiiiiii',   l'<-i> 

Till Iloiilal   solutions,  .",1 

Trypsin,    !;;•>,    fjs.  liiil 

artioii    of,    4K!i 
Trvpsiiioyi'ti.    tL'iJ,    t'_'s 
Tryplo|,liaiio,   .'.'.IL',   :>'.»>.    •'•I'l,   •'I'l- 

aml   yrowlli,  ."7ii,  ■"i7'< 
Tiiliiilis,  uriiiifcroiis,  fiiiiition   of,  ,"il7 
Tiiniors  anil  ilii't,  '>>*- 
Tiiiliiilitv  of  loiloi.ls,  ."il 
Tiiiik's'initlioil,    11.'.,  «;•-' 
Tyioi.'ill    pliiMioiiioioii,   lolloiils,   ."1 
Tyro.li-s  solution,    UN 
Tyrosino,   ."ni!.   lint,   li:'.!',   •"iiili 
Tiyptic-   .li>;rstioii.    pro'liic-ti   of,    lim 


I' 


I'ltraniiirosiopo,    Mini 
rni'ijnipi'iiMiti'il   aiiilosiM,  :t!t 
rnionililioni'il    nlli'x,    t:tl,   H.'ii! 
rnili-tiTiiiiiii'il    hitnufrii,  (illi,   ()29 
riolulatorv    movciiii'iit    of    rilis,   :il' 
Trra,  .■.'.'7,'  tins 
ill   Mooii,  I'llO 

iliiriiic   iliscaMi",   •i."'l 
I'Xiri'tion    of,   til.") 
rii'aso,  H'2,  fill! 
I'lii'  aciil,  ."-'!i,  ."1:11,  t>14,  (ilS 

amount    of,   ."iL'l' 

liasi's  of,  ."ill 

cliomii'ul  nature  of,  (i;t4 

I'liiloKi-noii!!  I'XiTction,  Ii47 

in   iliawi.si',   (iol 

metabolism  of,  fi4H 

of  blood,  648 

syntliesis  of,   644 

iimlir  ilruK**!  648 
I'rii'  aciil  iliatliesis,  l!.t4 


IMlKX 


rMi:{ 


rrii'fi*',    'ill' 
I'rii'i-iiiiii.    tl.'ii 
I'ririilttir    IikIix,   illl 
TriiH' : 

lin,!-  tit',   .'.ill 

iiiiiiiici  mill,  ''i.'lii 

iiriitiiatii'   iixviiriil-i   nf,   .".in 

lliloriilrn  . if,'. 'lid 

i'iiiii|iiiKitiiin,   ."i;i 

iTc'ilinliii'   iif,   ."iL".t 

il|.|in"<-iiiii   cif   fiti'/inn    |Miiiit   nf.    '-■'< 

lli|i|illlii-    liriil,    .".'(li 

liiiiiiiit,'i.|itiili'   iii'iil,   ."ill 

itKii'Uiinii-  riiiiMtitiii'iils  nt',  .';;i 

iiitnit^'rniiiii   (•«iiistitui.|it'*   nf.   .'iL*.! 

iHiriii.'il   iir^MMii'   s.'tltM   of,   .'ij.'l 

litiiisiiliiilrs,  :,:\2 

|i|iv-<ii'iil    |ir<ii'i"<si.>    iiiviihi.'l     ill    liimlii 
ti if.    I  I 

imriiir  liiiilir""  cif,  ."^".t 

nili'  iif  ■•vi-ri'licin.  lU.', 

rcai'tiiiri   t<f,    '>'2\ 

xk.'iliilv,    .V.I 

snliil    iiin»titlh.|itH    (if,    .'•_'." 

Nprritii-  yr.'ivity   nf,  .'C'L' 

»lll|illlltri   nf,    ."..lU 

t.il:il   |int..|itiiil  a.i.lity   nf,  .".-.'I 

uri':t   nf,  .'il-'T 

riillifiTnlis    llllillli.,    ."ill7 
rrnliiiillnm'ti,     I'.Mi 

rtili/atiiin   limit,  ti.'i:; 


Viiiiii'',    S7K 

.iirilriil   nf  Iicait,  L'17 

iiTi[iiil<i-i,  aflnrnit,  -1'- 
VayiH  ic'iitcr,  I'tTnct   nf   iiiintiiic  (III,   -^M 

ln...'ltii.li     nf,     L'L'L' 

tiinirity   cif,   L':.'! 
\:iiri|.<   iicrvr,   iiilliii'iiri'  nil   ri'>|.irat inn,   :\'A'2 
\aliii..,  tliM,  liciii 

Xiilvi's,   i.;irclia.',   ini'iliaiiisni    nf,    I"! 
aiirii.|iln\<.titrii'iit;ir,    l.'i  1 
iriniliinar,    l.'.l 
V:ill    Slykl'    lllftlln.l    fni-   a,.i,ln>is,    ILV    U 
Van    Slykr   inntliml    fnr   aniiim   ;;rnu|is,   (inn 
\  aNiiilar  n-lli'X,  I'sii 
\'arirn«'   viiiin,   "Jl  t 
Vasn.'nri.*triitinn.   'J'_'!l 
\'a.«ni-nnstrirtnr    lili-T^,    L*L'!' 

nii'tliii.ls  (if   ili'tri'tini;,   :.'■.",• 

nf  rxtri'initii's.  L'.'!.l 

nf   hia.l,    -M.I 

nf    visi-rra.    L'it.'l 

iiriuiii   nf,    L'.'li; 
Vasnililatiir    fillers.    1.':I4 

iiii'tliiiih    fnr   ilcli'itiiit;,   L':.'!! 

nrit'in  nf.   -.It 
Vasnililatiir   ru'rvrs,  7!'7 
Vasnrniitnr   ccafiT: 

nffiTPnt  iinjMilsi's.  L':!X,  2:i!1 

cliii'f   ci'iitiT,   LI.'!."i 

fffi'i't   nf   JI  inn   (if  liliin.l   nn,  '2^H 


VaHiinmtni  (.niter   -  <  *t.nt  'il 

li(trin(ini>   ediitrnl    nf,   -.'17 

•dilixi'liary  centers,   '-M"i 
VaHnnintiir   tiliers,   J.'ll 

(irit;in  nf,  L'ilJ 
Vnsdtnnic   iniinilses,  -In 
Veins,   ili^a|i|iearani'e   nf   pulse   in,   '2i''> 
Velniity   rniistant,   enzymes,   7" 
Velncity,  mean   lineal,  L'nti 

pulse,   ::iMP 
Venniis   lilnnil,  tcnsinn   nf  •'(»    in.  .';|'J 

Vellnlis    nlllllllW,     U:in 

N'enniis   pulse  trai'in^.  1'7.'( 

Vilnius  return  tn  I.e.irt,  L'!''-' 

X'euniis  sinus.  LMS 

Veutllatiiin  nf  Iniiijs,  .'t.'ii 

\  eiilriile,  (.nrv(  s   nf   pressure   in,    1  Iti.    IIH, 

|."il 
Ventricles: 

cnncluc-tivity     tissue     itt\     IHi 

tilirillatinii,"  IK.'i 

spreail  nf  lie.it   in.   )'■<:.   I!U 
Viirnin.   ."i7S 

\  isc.ira.   111 1   suppiv    nf,   ■:\7 

Visceral   111 llIdH,  j'lL' 

VisciTal    seiisiliveness,    SS.'i 
Viscnsity    nf    111 1,    nil 

Visual   (■(. liter,  x.'il 

\'isu:il    psycliic   areas,   s",  ( 

\'i>n:il    s(. usury    arc.;!,    H.'.  | 

Vital   activity',    1  » 

Vital  capacity,  linn,   :;l.". 

in  ilisc.'isi.,  .':i  \ 
\'it:(l   tli(.nry   nf    urine   e\c.refinn,   .*i7L* 
Vitaniiiies.  .'H\ 

Vividiffiisi iliMl 

N'nlitinll,    7'<li 
Vciniitiiii;,    I  !!• 

W 

Water   cc.Mteiit    nf   M 1,   sd 

Wati  r    lianinicr, 

ill   lilc.ci.l   pressure   measure. inent.   1.'!:; 
Wlieatstnni.   Iiridye,    IS 
White   cres.-elitic    line,    -L'ti 
Wi^i;ers   ni.'iiiniiieter,    1  Itl 
Willard   (iil.l.s,   principle   „(,  M 
Wnr.l    liliu.liicss.    sc,:: 
Wnrd    cc.lltcrs.    Ml:; 
Wnr.l    c|eafnes«.   Hti;; 

X 

X.Tiitliine,   liii.l 

Xanthine   cixidase,   li:!!! 

\;inllic.sine.  (i:!li 

X  rays,  in  study  nf  stcimaeli,  4.'i;i 

iiinveineiits   nf   stomach   seen    liv   aid   (if, 
451 

Z 

Zein.   inadec|iiacy   fnr   ({rciwtli,  ."i7.S 
Zyninyen    yraiiules.    IL'o,   4L.'l,    t2!t 


